t ' ANNALS OF THE MISSOURI BOTANICAL GARDEN Annals of the Missouri Botanical Garden Volume 1933 With Twenty-three Plates and Forty-three Figures Published quarterly at 8 West King Street, Lancaster, Pa., by the Board of Trustees of the Missouri Botanical Garden, St. Louis, Mo. K Entered as second-class matter at the post-office at Lancaster, Pennsylvania, under the Act of March 3, 1879. 5 1306 Annals of the Missouri Botanical Garden A Quarterly Journal containing Scientific Contributions from the Missouri Botanical Garden and the Graduate Labora- tory of the Henry Shaw School of Botany of Washington Uni- versity in affiliation with the Missouri Botanical Garden. Information times the calendar year: February, April, September, and November. Four numbers constitute a volume. Subscription Price $6.00 per volume Single Numbers 1.50 each Contents of previous issues of the Annals of the Missouri Botanical Garden are listed in the Agricultural Index, published bv the H, W. Wilson Co. * v I STAFF OF THE MISSOURI BOTANICAL GARDEN Director, Geor<;e T. Moore. Assistant to the Director, Katherine H. Leigh, Hermann von Schrenk, Pathologist. Jesse M. Greenman, Curator of the Herbarium Ernest S. Reynolds, Physiologist. Carroll W. Dodge, Mycologist. Robert E. Woodson, Jr., Research Assistant. Nell C. Horner, Librarian and Editor of Publications. BOARD of trustees OF THE MISSOURI BOTANICAL GARDEN President, George C. Hitchcock Vice-President, Samuel C. Davis. Second Vice-President, Daniel K, Catlin. L. Ray Carter. Thomas S. Maffitt George T. Moore. Albert T. Perkins. Eugene Pettus. Ethan A. H. Shepley Fred G. Zeibig. EX-OFFICIO MEMBERS: George R. Throop, Chancellor of Washington University. William Scarlett, Bishop of the Diocese of Missouri. Bernard F. Dickmann, Mayor of the City of St. Louis. Albert Kuntz, President of The Academy of Sci ence of St. Louis. David C. Todd President of the Board of Education of St. Louis. George F. Heffernan, Secretary. TABLE OF CONTENTS PAGE A New Yellow Nymphaea from Tropical Africa George H. Pring and Robert E. Woodson, Jr 1-ti Contribution to the Lichen Flora of North America Veli J. P. B. Rasanen 7-21 Astranthium and Related Genera Esther L. Larsen 23-44 Ferns and Fern Allies of Missouri M. Elizabeth Pinkerton 45-78 Blastomycosis : Report of a Case, with a Study of an Etiologic Factor and a Classification of the Organism Morris Moore 79-118 A Monograph of the American Species of the Genus Halenia Caroline K. Allen 1 19-222 The Sensitivity of Orchid Seedlings to Nutri- tional Ions F. L. Wynd 223-237 A Revision of the North American Species of Verbena Lily M. Perry 239-362 Nutrient Solutions for Orchids F. Lyle Wynd 363-372 The Foliose and Fruticose Lichens of Costa Rica. I Carroll W. Dodge 373-467 A Neutral (?) Strain of Mucor sphaerosporus from Missouri Morris Moore 469 A Study of Endomyces capsuUitus Rewbridge, Dodge and Ayers: A Causative Agent of Fatal Cerebrospinal Meningitis Morris Moore 471-568 Sources of Carbohydrate for Germination and Growth of Orchid Seedlings F. Lyle Wynd . . 569-581 PAGE Super Optimal and Thermal Death Tempera- tures of the Cotton Plant as Affected by Variations in Relative; Humidity .... Dorothy Megowen Berkley and Earl E. Berkley 583-004 Studies in the Apocynaceae. IV. The Amer- ican ( loner a of Echitoideae . . Robert E. Woodson, Jr. 005-790 Report of a Botanical Expedition into the Mountains of Western Texas Julian A. Steyermark and John Adam Moore 791-800 General Index to Volume XX 807-817 Annals of the Missouri Botanical Garden Vol. 20 FEBRUARY, 1933 No. 1 A NEW YELLOW N YMPHAEA FROM TROPICAL AFRICA GEORGE H. PRING Superintendent j Missouri Botanical Garden ROBERT E. WOODSON, Jr. Research Assistant, Missouri Botanical Garden on Nymphaea (§ Brachyceras) Burttii Pring & Woodson, spec, no v., foliis magnis longe-petiolatis orbiculare-sagittatis margine plus minusve conspicue undulato-sinuatis apice late obtusis vel rotundatis sinu profundo auriculis leviter divergentibus obtusis 25-35 cm. longis subcoriaceis utrinque viridibus vel saepius purpureo-maculatis glaberrimis vel umbilico paulo papillato subtus nervis manifestis sed vix prominentibus ; floribus speci- osissimis primulino-flavis 15-18 cm. diametro metientibus, sepalis ovato-lanceolatis acutiusculis 5-8 cm. longis 2-3 cm. latis dilute viridibus immaculatis laevibus, petalis ca. 20-23 anguste ellipticis apice acuminatis vel anguste acutis medio versus grada- tim angustatis plerisque 5-nervatis exterioribus quam sepalis paulo brevioribus ca. 4.5-7.0 cm. longis 1.0-2.5 cm. latis, stami- nibus 190-200 linearibus connectivo manifeste elongato basi paulo ampliato exterioribus ca. 5 cm. longis dilute cadmio-flavis, carpellis 28-30 stylo linearo profunde inclinato stigmate concavo ; fructu minore 3.5-4.0 cm. diametro metiente, seminibus ovoideo- oblongoideis apice minutissime apiculatis longitudine leviter punc- ticulato-striatis griseo-brunneis maximo ca. 0.075 cm. minimo ca. 0.05 cm. diametro metientibus. — Cult. Missouri Botanical Garden, Aug., 1930, G. H. Pring s. n. (Herb. Mo. Bot. Garden, type) . Leaves large, long-petiolate, orbicular-sagittate, margin more or less conspicuously undulate-sinuate, apex broadly obtuse or rotund, sinus relatively deep and narrow, auricles obtuse, slightly 1 Issued April 29, 1933. Ann. Mo. Bot. Gard., Vol. 20, 1933. (1) [Vol. 20 2 ANNALS OF THE MISSOURI BOTANICAL GARDEN divergent, 25-35 cm. long, subcoriaceous, either surface green, or occasionally somewhat purplish-maculate, particularly when young, glabrous, or the umbilicus slightly papillate, the nerves manifest but not prominent beneath; flowers showy, primrose- yellow, veiy fragrant, 15-18 cm. in diameter; sepals ovate- lanceolate, acute, 5-8 cm. long, 2-3 cm. broad, pale green, without purple spots, smooth; petals about 20-23, narrowly elliptic, apex S. S.L. P P. L. Woodson S.I<., submerged leaves; F. L. t floating leaves. acuminate to narrowly acute, the base gradually narrowed from about the middle, 5-nerved, the exterior somewhat shorter than the sepals, usually 4.5-7.0 cm. long, 1.0-2.5 cm. broad; stamens 190-200, linear, the exterior about 5 cm. long, cadmium-yellow, the connective manifestly elongate, somewhat broadened at the base; carpels 28-30, the style linear, sharply inclined, the stigma concave; fruit relatively small, about 3.5-4.0 cm. in diameter; seeds ovoid-oblongoid, minutely apiculate, rather inconspicuously 1933] PRING AND WOODSON — A NEW YELLOW NYMPHAEA 3 puncticulate-striate longitudinally, about 0.075 cm. long, 0.05 cm. broad, grayish-brown. — Cultivated at the Missouri Botanical Garden, Aug., 1930, G. H. Pring s. n. (Herb. Mo. Bot. Garden, type) . Only two yellow tropical Nymphaeas have previously been described from Africa: N. sulphurea Gilg and N. Stuhlmannii Schwfth. & Gilg. The former differs from N. Burttii chiefly in the smaller flowers (4-7 cm. in diam.) and leaves (4.5-5.5 cm. long), which are almost exactly orbicular-cordate in outline, purplish-maculate sepals, and more elongate tubers. N. Stuhl- mannii is easily distinguishable from N. Burttii by the somewhat smaller flowers (10-15 cm. in diam.), with broader, obovate, obtuse or rounded petals, and smaller (21-25 cm. long), entire leaves, which are broadly ovate-cordate in outline, with broad, rounded, regular lobes. Furthermore, the venation of the leaves of N. Stuhlmannii is extremely verrucose beneath. The type specimen of N. Stuhlmannii has not been available to the writers for personal examination, but has been compared with a duplicate specimen of N. Burttii by Dr. H. Melchior, of the Botanical Museum at Berlin-Dahlem, who kindly affirmed the distinction of either species. A photograph of the type specimen of N. Stuhlmannii (Stuhlmann 1^10 in Hb. Berol.), generously provided by Dr. L. Diels, Director of the Botanical Garden and Museum at Berlin-Dahlem, has been deposited in the herbarium of the Missouri Botanical Garden. After a search of over ten years, the director of the Missouri Botanical Garden obtained in September, 1929, a seed-pod of what was presumed to be Nymphaea Stuhlmannii through the personal efforts of Mr. B. D. Burtt, Esq., botanist for the Tsetse Research Bureau, Kondoa, Tanganyika Territory. An excerpt from Mr. Burtt's notes is quoted. "The seed was collected from plants growing in a seasonal rain-pond in the Sambala ' Mbuga' seasonal swamp, the plants having spectacular yellow flowers 8 inches in diameter and sweet scented. The flowers float on the surface of the water and on examination were found to contain dead bees (Apis melifera I?]) that were imprisoned by the anthers over the stigmatic surface of the flowers. The plants were collected on May 19, 1929, the seed from the same locality on July 15, 1929. Other plants were collected March 16, 1929, from a seasonal rain pond near Salia, Kondoa Distr. I have observed the plant in seasonal rain ponds at Magungila, Wembare Steppe in 1928, also near Lilbilin, Massai Land, in 1927." [Vol.. 20, 1933] 4 ANNALS OF THE MISSOURI BOTANICAL GARDEN The single, small seed-pod, in somewhat immature condition, was received at the Garden on September 19, 1929. It was im- mediately cleaned, and the many seeds planted in the greenhouse water-lily tanks. Three weeks later a single seed germinated from the lot, producing its first primrose-yellow flower on June 17, 1930. On July 30, the plant was removed from its pot and plant- ed in the pond out of doors where it continued to bloom until frost. The successful introduction of a yellow tropical water-lily has opened an entirely new field for the hybridist. For many years the available colors in the Brachyceras group have been limited to blue and pink, and more recently to white through the intro- duction from the Missouri Botanical Garden of the hybrid "Mrs. G. H. Pring." During the past three seasons 250 pol- linations have been made with the pollen of N. Burttii, resulting in many variable forms of commercial interest. The first season (1930) was devoted to selfing the yellow-flowered species, and many fertile seed-pods resulted. The following year twenty specimens were grown from seed and planted in the ponds out of doors. No variation in the color of the flowers was observed, but a variable character was evident in the peduncle and petioles, some being pure green, whileabout an equal number were brownish. An unusual character, which appears to be dominant in all the plants, is a peculiar twisting of the peduncle, the day before the flower opens, in such a manner as to submerge the bud. The following morning, however, the peduncle straightens, holding the open flower erect in a natural position. This action has also been observed in several hybrids. As a propagator, Nymphaea Burttii is by far the poorest of any grown at the Garden. Of the fifty propagating tubers secured from pot plants during the past two seasons, only one has pro- duced growth up to the present, despite the fact that they have been in the heated propagating tanks for a period of six months. Since it is an extremely poor propagator from tubers and extreme heat is necessary for growth, it is very doubtful if it will find a place in garden pools. On the other hand, the hybrids derived from it, particularly the "Saint Louis," are both readily propa- gated from tubers and suitable for cooler water. [Vol. 20, 1933] 6 ANNALS OF THE MISSOURI BOTANICAL GARDEN Explanation of Plate PLATE 1 Leaf, bud, and flower of Nymphaea Burttii Pring & Woodson, X %. del. A. A. Heinze Ann. Mo. Hot. Gard., Vol. 20, L933 l'LA 1 l'KINO & WOODSON— A MOW YELLOW NVMI'll AKA CONTRIBUTION TO THE LICHEN FLORA OF NORTH AMERICA^ VELI J. P. B. RASANEN Instructor in the Agricultural School, Kurkijoki, Finnland The lichens here described have been collected in the northern section of North America. Of these specimens, 18 are Alaskan, 60 from eastern Canada (New Brunswick, except one from Quebec), approximately 150 from western Canada (British Columbia and Alberta), and a few from the state of Wyoming. The Alaskan lichens, 15 species in all, are largely the usual arctic ground lichens. The New Brunswick lichens, which com- prise 46 species, come from around Dalhousie and were collected in the year 1930 by the agronomist Tapio Reijonen. Usnea longissima was collected near Franklin, Quebec, by Dr. Viljo Kujala in 1931 during his scientific journey. On this same journey, he collected in western Canada (120 species, varieties, and forms) and in Wyoming (6 species). The latter specimens come from the sandstone in a cactus desert. The lichens from western Canada were collected by Dr. Viljo Kujala in various types of forest in British Columbia in the following localities: Jackman, Longworth, Aleza Lake, Prince George, Six Mile Lake (Trout Lake), Hazleton, Dorreen, Co- wichan Lake (Vancouver Island), Kamloops, Fish Lake (near Kamloops), Blue River, Valemount, Golden, Field, and in Al- berta in the vicinity of Lake Louise. The desert lichens come principally from the vicinity of Kamloops in British Columbia and from Wainwright in Alberta. Although the same species have been encountered in several different localities, the total number of species, varieties, and forms is 171 with 5 entirely new species, as follows: Usnea Kujalae, Placodium xanthostigmoideum , Nephromium canadense, Diplo- schistes canadensis, and Sphaerophorus Tuckermanii. The list further includes eleven new varieties and forms, as follows: Usnea comosa var. stuppea, Alectoria sarmentosa var. gigantea, A . Fremontii f . perfertilis, Cetraria nigricans var. pallida, 1 Duplicates of most of the specimens have been deposited in the herbarium of the Missouri Botanical Garden. Issued April 29, 1933. Ann. Mo. Bot. Gard., Vol. 20, 1933. (7) 8 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 20 C. tenui folia var. reticulata, C. tenuifolia var, pseudoislandica f. septentrionalis, C. juniperina var. canadensis, C. juniperina var. crispata, Parmelia elegantula var. americana, Cyanisticta Hookeri var. septentrionalis, Myeoblastus sanguinarius var. Dodgeanus. Species new to North America or otherwise noteworthy include the following: Usnea lapponica, U. dasypoga, U. caucasica, U. prostrata, U. rugulosa, U. similis, Alectoria Fremontii subsp. olivacea, A. jubata var. Vrangiana, Ramalina Roesleri, Ochrolechia upmliensis, Pertusaria xanihostoma, Physcia pyrithrocardia, P. muscigena f. squarrosa, P. endophoi nicea, Peltigera Maurizii, P. canina var. suomensis, P. lepidophora, P. Nylanderi, Cetraria Merrill ii, Parmelia Delavayi, Squamaria alphoplaca, Nephromium subparile, Lobaria oregana, Diploschistes bryophiloides. The lichens collected by Dr. Kujala are principally epiphytic and soil-lichens from a variety of forest types, although a few of the soil lichens come from fields or desert. It might be inter- esting to note here that in regions as widely separated and isolated as the North American prairies and the south Russian and Asiatic steppes and deserts there appear many identical, or, at least similar, lichen forms living on the ground. Such steppe lichens are: Parmelia vagans (appearing also in south Russia and Asia), Physcia muscigena (many forms, a few also growing in the mountains), Acarospora Schleicheri (appeal's in the Asiatic, North- African, and Russian steppes, in addition to the European Alps and Pyrenees), 1 Psora decipiens (also in deserts and mountains), Diploschistes canadensis (the similar Russian D. scruposus var. terrestris), Ochrolechia upsaliensis (sporadically found outside of the steppes), Cladonia cariosa var. cribrosa (particularly on other mineral substrates, slightly calciphile). As the lichens in this collection were not collected by a lichen- ologist, the lichen flora of the various localities is very poorly represented and poor in species. They are, however, of particular interest to the European investigator on account of the many similar forms. The typical European Cetraria juniperina and C. islandiea do not appear at all amongst the American lichen specimens (the latter also being absent from the u Cetraria islandiea" specimens collected in Tierra del Fuego by Professor Auer's expedition in 1929). i Magnusson, Monogr. Acarospora. p. 395, 1929. 1933] RASANEN CONTRIBUTION TO LICHEN FLORA OF AMERICA 9 As the lichens from the various localities have not been suffi- ciently collected, conclusions as to the relationship between the American and European species are still hypothetical, particularly as the present systematic revision of lichens necessitates many corrections of the earlier determinations. We can, however, state that as far as our present knowledge extends a great number of identical lichen forms appear both in North America and Eurasia, although the quantitative relationships between the different species differ considerably on the two sides of the Atlantic. I wish to acknowledge with thanks Dr. C. W. Dodge, My- cologist to the Missouri Botanical Garden, who has kindly trans- lated my manuscript from German to English, and T. Hiden, of Helsinski, Finnland, who has gone over the latin text of this work. Usnea DA8TPOGA (Ach.) Rohl. New Brunswick: Dalhousie; British Columbia: Prince George and Six Mile Lake, on Picea. Sterile. Usnea dasypoga var. subscabrata Vain. British Columbia: Hazleton, on Betvla and Pinics. Sterile. Usnea caucasica Vain. Thallus prostratus, glauco-viridis, medulla sublaxa, K— . — British Columbia: Six Mile Lake, on Picea. Fertile. Usnea prostrata (Vain.) Ras. British Columbia: Aleza Lake, on Picea. Sterile. Usnea rugulosa Vain. British Columbia: Hazleton, on trunks of Pinus Murray- ana. Sterile. Usnea longissima Ach. Quebec: north shore of the St. Lawrence Bay, Franklin, especially on branches of Picea in a mixed forest; British Columbia: Vancouver, Catillano Canyon, on Thuja in a forest with abundant ferns. Sterile. Usnea comosa (Ach.) Vain. Thallus erectus, parce sorediosus, soredia isidiosa; medulla crebra, K — . — New Brunswick: Dalhousie, on Picea; British Columbia: Hazleton, rarely on branches of Pinus Murrayana. Sterile. Usnea comosa var. stuppea Ras., var. nov. Thallus erectus aut suberectus, brevior fruticulosus, 5-7 cm. longus, laevigatus vel leviter verrucosus, sorediosus, pallido-stramineus; soredia maculiformia, demum parce isidiosa. Medulla laxa, stuppea, K — . — British Columbia: Hazleton, very frequent on small, half -dried Pinus } Picea y and Betula, on a sandy heath. Sterile. Usnea similis Motyka U. subcomosa Vain, in Kgl. Danske Vidensk. Selsk. Skrifter, Nat. og Math. Afd. VIII. 6:392 (110). 1924. With the above species, occasional in Hazleton. Sterile. Usnea lapponica Vain. With the above species, Hazleton, on Pinus. Sterile. Usnea hirta (L.) Motyka. British Columbia: Aleza Lake, occasional on Picea canadensis and P. Engelmannii. Sterile. Usnea Kujalae 1 Ras., sp. nov. Thallus curtus, erectus, fruticulosus, 3-5 cm. longus, bene ramosus, lacteo-stramineus vel albo-viridis, sorediosus. Rami laevigati, sine verruculis nervisque. Medulla laxissima, K — . Apothecia rara, terminalia, 2-3 mm. lata, albido-straminea, fibrillosa, plana. Sporae 9-10 X 5.3-7 m, ovoideo- 1 In honor of Dr. Viljo Kujala. 10 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 20 ellipsoidene, 8 : nae, monostichae. Asci cylindrici. Hymenium 80 /x crassum, I+, caerulescens. Subsimilis Usnem sorediiferue Motyka, sed thallus albidus et parce sorediosus. — British Columbia: Hazleton, abundant on branches of Pinus Murray- ana. Rarely fertile. Alectoria sakmentosa Ach. Thallus elongatus, pendulus vel prostratus, K = K(C1) =F leviter roseus, demum ferrugineus, I=F leviter caerulescens. Sporae normaliter binae vel 4: nae, fuscae. — British Columbia: Aleza Lake, frequent on branches of Abies lasiocarpa, Picea canaxU nsis, and P. Engelmannii; Longworth, frequent on branches of Thuja; Hazleton, rarely on Pinus. Fertile. Alectoria sakmentosa var. gigantea Riis., var. nov. Thallus elongatus, pendu- lus vel prostratus, 50-80 cm. longus, albido-stramineus, parce pseudocyphellatus, esorediatus, K = , K(C1)=, Cl = , 1= leviter caerulescens. Apothecia vulgaria, plana vel concava, atro-fusca vel livido-fusca, nuda vel leviter pruinosa. Sporae normaliter 3: nae, 25-31 X 14.5-20 m, ellipsoideae, fusco-mgrae. Hymenium 130 m crassum, 1+ caerulescens. — British Columbia: Longworth, frequent on Thuja; Dorreen, on Pseudotsuga. Alectoria laeta (Tayl.) Linds. A.japonica Tuck., A. osteina Nyl., A. lata DR. (Arkiv f. Bot. 20 u : 24. 1926). Thallus suberectus vel prostratus, osteo-ochroleucus, apicem versus concolor, Kzfc passim obsolete intensive fulvescens, K(C1) T roseus. — British Columbia: Hazleton, occasional on the ground ; Alaska, occasional on the ground. Sterile. Alectouia ochroleuca (Ehrh.) Nyl. Thallus pro parte ochroleucus, majore parte vindi-niger, K= vel basim versus obsolete =k dilute fulvescens, K(C1) = Alaska, on the ground. Sterile. Alectouia implexa (HofTm.) Nyl. f. fuscidula Arn. Thallus pallido-fuscescens, K± lutesceni, sorediosus. Soredia albida. — British Columbia: Hazleton, on branches of Pinus, Aleza Lake, on branches of Abies and Picea. Sterile. Alectoria jubata (L.) Nyl. var. prolixa Ach. Thallus fuscescens vel nigro- fuscescens, K = , sorediosus. Soredia copiosa, albida. — New Brunswick: Dalhousie, on trunks of Abies; British Columbia: Kamloops, Fish Lake, Golden, and Aleza Lake, very frequent on branches and trunks of Pseudotsuga and Pinus Murrayana. Also on conifers near Hazleton. Sterile. Alectouia jubata var. Vrangiana (Gyeln.) Ras. A. Vrangiana Gyeln. (Magyar Bot. Lap. 31: 46. 1032). Thallus parce sorediosus, vel fere esorediatus, olivaceo-viridis vel fusco-viridis, K=, K(C1)=. — British Columbia: Hazleton, on conifers, occasional. Sterile. Alectouia Fkkmontii Tuck. f. perfertilis Ras., forma nov. Thallus prostratus, obscuro-castaneo-fuscus, laevis, subnitidus, esorediatus, K = , Cl=, K(C1)=. Apothecia vulgaria numerosissima, lateralia, convexa, flavida. Sporae 5-8 X 5 m, ovoideae, 8 : nae, hv.ilinae. Hymenium 1+ caerulescens. — British Columbia: Golden, on the trunks and branches of Pinus Murrayana. Alectouia Fuemontii subsp. olivacea Ras. Thallus olivaceo-castaneus, prostratus, laevis, nitidus, esorediatus, K = , Cl=, K(C1)=. Apothecia rarissima, minora, flavida. — British Columbia: Golden, on Pinus Murrayana and Pseudotsuga, Kamloops, Fish Lake, on Pinus Murrayana. Sterile. The sorediose form, named by Du Rietz (Arkiv f. Bot. 20 u : 8. 1926) subsp. Eriks- soniij was not seen among the North American specimens. It would appear more correct, as Du Rietz (Z. c.) thinks, not to regard the sorediose form as the main species. My subspecies olivacea (Riisanen, Medd. Soc. Fauna Flora Fennica 43: 4. 1933] RASANEN AMERICA 1 1 typ so that we have here three different Alectoria types. Ramalina thrausta (Ach.) Nyl. British Columbia: Hazleton, on branches of conifers, Prince George and Six Mile Lake, on Picea. Sterile. Ramalina geniculata Nyl. British Columbia: Hazleton, on branches of Pinus Murrayana, rare. Sterile. Ramalina roesleri (Hochst.) Nyl. New Brunswick: Dalhousie, on branches of Picea, Sorbus, and on a wooden wall. Sterile. Ramalina calicaris (L.) Fr. New Brunswick: Dalhousie, on trunks of Populus. Fertile. VULPINA Thallus sorediosus. Medulla 1+ caerulescens. British Columbia: Kamloops, Fish Lake, and Field (subalpine), on branches and trunks of Pseudotsuga. Sterile. Letharia vulpina f. incompta Ach. Thallus sorediosus et bene isidiosus, me- dulla 1+ caerulescens. — Found with the species near Fish Lake, Golden, and Jackman, B. C, on Pinus Murrayana. Sterile. Cornicularia divergens Ach. Alaska (alpine), on the ground. Sterile. Cetraria Merrillii DR. C. californica var. Tuckermanii Howe Thallus 0.5-2 cm. longus, rigidus, erectus, divaricato-ramosus, opacus, olivaceo- niger. Rami angulato vulgaria, 0.5-3 plana vel convexa, terminalia, atra vel fusco-nigra, nuda, subnitida, subciliata. Sporae 8:nae, 6-8 X 4-5.5 m, ovoideae, incoloratae. — British Columbia: Golden, Kamloops, Fish Lake, and Hazleton, on branches of Pinus Murrayana and other conifers. Fertile. Cetraria nigricans Nyl. var. pallida Ras., var. nov. Thallus superne pallidus vel olivaceo-pallidus, subtus albidus, basim versus sanguineo-lentus; medulla 1 + caerulescens. Laciniae 1-2 mm. latae, subcanaliculatae, denticulato-ciliatae, dicho- tome ramosae. — Alaska, on the ground with Alectoria ochroleuca and Cornicularia divergens. Sterile. Cetraria tenuifolia (L.) Vain. Thallus canaliculars, laevigatus, medulla K — . — Alaska, rarely with the previous species. Sterile. Cetraria Thallus subtus foveato- reticulatus, castaneo-fuscus, in marginibus dense isidioso-dentatus. Laciniae angustae, canaliculato-tubulosae, crispatae, marginem versus pseudocyphellatae. Medulla K — . — British Columbia: Kamloops, on stones, Valemount and Aleza Lake, on the ground. Sterile. Cetraria tenuifolia var. pseudoislandica Ras. (Ann. Bot. Soc. Zool.-bot. Fenn. Vanamo 2: 15. 1932) f. septentrionalis Ras., forma nov. Medulla I-. Thallus similis C. islandicae, sed I-, et margo laciniarum fere sine spinis vel ciliis. Color pallido-castaneus. — British Columbia: Golden, on trunks and branches of dwarf shrubs near the ground. Sterile. The true Cetraria islandica (L.) Ach., which gives the reaction "Medulla I lutes- cens, demum ferrugineo-rubescens," has not been seen by me from either North or South America. Cetraria nivalis (L.) Ach. British Columbia: Kamloops, on the ground and rocks in semi-desert; Alaska, on the ground. Sterile. Cetraria cucullata (Bell.) Ach. British Columbia: Valemount, on the ground with Sterile. [Vol. 20 12 ANNALS OF THE MISSOURI BOTANICAL GARDEN Cetkauia caperata (L.) Vain. C. pinastri (Scop.) Rohl. British Columbia: Hazleton, a small specimen with other Cetraria species on branches of Pinus Murrayana, Aleza Lake, a chiefly geoplese lichen (Riisiinen, t)ber Flechtenstand u. Fl. Veget. im Westl. Nordfinnl., p. 84. 1927), with Parmelia ambigua and P. pallescens on the base of Abies lasiocarpa. Sterile. Cethakia juniperina (L.) Fr. var. viridis (Schweinitz) Ras. C. viridis Schweinitz apud Halsey, Ann. Lyceum Nat. Hist. New York 1: 16. 1824; C. juniperina var. virescens Tuck. Thallui curtus, rigidus; laciniae obtusae, obscuro-virides; K = , Cl=, I=. Apothecia terminalia, majora, 3-8 mm. lata, fusco-nigra, subnitida, margo sub- ereaata, tenuis. Sporae 8 : dm, ovoideae, 6-8 X 3.5-6.5 m, incoloratae. — British Columbia: Golden, common on branches of Pinus Murrayana. Fertile. Cetraiua juniperina var. canadensis Rils., var. nov. Thallus erectus vel sub- erectus, subrigidus; laciniae obtusae, aureo-flavae vel interdum viridi-flavae, super retirulato-nervosae, intus flavar. Apothecia terminalia, minora, 1-3 mm. lata, castaneo-fusca, nitida; margo lacerato-crenata. Sporae 6.5-8 X 4-5.2 », ovoideae, 8:nae, hyalinae. — British Columbia: Kamloops, Fish Lake, Aleza Lake, and Hazleton, very frequent on twigs of conifers. C. juniperina (the main species) has apothecia chiefly in the middle of the thallus and the lobes are very divided. Also, the color of this species is not so bright a golden yellow as in the variety canadensis, and the mirface is furthermore much smoother in the species. Cetraiua juniperina var. crispata Run., var. nov. Sicut var. canadensis, sed thallus dense lacerato-crispatus vel subpapillatus. Apothecia rara. Est analogica cum C. juniperina var. terrcstris Schaer. (= C. Tilesii Ach. in Zahlbr. Cat. Lich. Univ. 6: 341. 1930). — British Columbia: Golden, rarely on branches of Pinus Murrayana with var. viridis. Fertile. Cetraiua aleurites (Ach.) Th. Fr. New Brunswick: Dalhousie, on trunks of Abies and on wood. Sterile. Cetraiua chrysantha Tuck. Plat ys ma septentrionale Nyl. Thallus stramineus, lacunoso-nervosus, esorediatus, subtus fusco-niger, K± lutescens, Cl=, K(C1)t roseus, I=. — Alaska (alpine), on the ground. Sterile. Cetraiua lacunosa Ach. var. Macounii DR. Thallus subtus fusco-niger, non isidiosus, I =. — British Columbia: Kamloops, on stones in semi desert. Sterile. Cetraiua norvegica (Lynge) DR. Thallus isidiosus, 1=.— With the above species in Kamloops, on stones. Sterile. Cetraria glauca (L.) Ach. Thallus I=F caerulescens. — With the above species in Kamloops, on stones; Prince George, Six Mile Lake, abundant on Picea; Aleza Lake and Vancouver, on branches of Picea. Sterile. Cetraria glauca var. stenophylla Tuck. (Syn. North Amer. Lich. 1 : 36. 1882). Thallus I T caerulescens, anguste laciniatus, glaucus, subtus niger. — British Columbia: Vancouver Is., Lake Cowichan, occasional on the lower branches of Picea in the forest. Sterile. Cetraria scutata (Wulf.) Poetsch C. chlorophylla (Willd.) Dalla Torre et Sarnth British Columbia: Kamloops, Fish Lake, Golden, Hazleton, Aleza Lzke, on the branches of Pinus Murrayana and Abies; Vancouver Is., Lake Cowichan, occasional on Thuja gigantea. Sterile. 1933] RASANEN — CONTRIBUTION TO LICHEN FLORA OF AMERICA 13 PLATYPHYLLA Cetraria platyphylla Tuck. Thallus 1-3 cm. latus, irregulariter laciniatus, super opacus, obscure fuscus, tuberculosus, subtus pallide fuscus, reticulato-nervosus. Apothecia marginalia, snata. Sporae minora crassum, 1+ caerulescens. 4-6.5 m. Hym Murray ana. Nephromopsis ciliaris (Ach.) Hue. British Columbia: Aleza Lake, a small specimen on Pinus Murrayana. Sterile. Parmelia pallescens (Neck.) Ras. P. hyperopta Ach. New Brunswick: Dalhousie, on trunk of Abies; British Columbia: Aleza Lake, on base of Abies lasiocarpa as a geoplese lichen. Sterile. Parmelia ambigua (Wulf.) Ach. New Brunswick: Dalhousie, on wood; British Columbia: Aleza Lake, on base of Abies lasiocarpa and Pinus Murrayana. Sterile. Parmelia physodes (L.) Ach. f. labrosa Ach. New Brunswick: Dalhousie, on trunk of Abies; British Columbia: Aleza Lake, on trunks of Picea t Kamloops, on Pinus Murrayana. Sterile. Parmelia physodes f. vittata Mereshk. British Columbia: Aleza Lake and Hazleton, on branches of Pinus Murrayana. Sterile. Parmelia lophyrea Ach. Thallus cinereus, esorediatus, K± lutescens, K(C1) roseus. Apothecia vulgaria, subtus ampullacea, demum ca. 5 mm. lata. Sporae sphaeroideae, 2.5-5 m, 8 : nae, incoloratae. Hymenium 50 /x crassum, 1 + caerulescens. — British Columbia: Golden, on branches of Pinus Murrayana; Aleza Lake, on branches of Picea. Parmelia delavayi (Hue) Nyl. Thallus parvus, esorediatus, obscure cinereus. Apothecia vulgaria, demum 5 mm. lata, subtus non ampullacea, discus fuscus, urceolatus vel deplanatus. Sporae 5-7 X 4-5 n f 8 : nae, ovoideae. Hymenium ca. 55 m crassum, 1+ caerulescens. — British Columbia: Golden, on Pinus Murray- ana. Parmelia enteromorpha Ach. Thallus esorediatus, ventricoso-inflatus, cavus, KzL lutescens, K(C1)=. — British Columbia: Vancouver Is., Lake Cowichan, on trunk of Thuja gigantea. Sterile. Parmelia vittata (Ach.) Rohl. British Columbia: Prince George, Six Mile Lake, on trunk of Betula?. Sterile. Parmelia sulcata Tayl. New Brunswick: Dalhousie, on trunks of Betula, Sorbus, Picea , Abies, and on wood; British Columbia: Prince George, Six Mile Lake, on Betula; Hazleton, on Betula; Golden, on Pinus Murrayana and Pseudotsuga; Kamloops on arid ground. Sterile. Parmelia vagans Nyl. P. molliuscula Tuck. (Syn. North Amer. Lich. 1: 64. 1882). Thallus dichotome laciniatus, sine sorediis et isidiis, stramineo-virens, K T f ulves- cens. — British Columbia: Kamloops, on the ground between grasses and shrubs. Sterile. The lichen is a typical desert lichen and appears also in such environment in Russia and Asia. Parmelia subaurifera Nyl. unk trunk of a Populus. Sterile. Parmelia papulosa ( P. exasperatula Nyl. [Vol. 20 14 ANNALS OF THE MISSOURI BOTANICAL GARDEN New I trims wick: Dalhousie, on the trunk of Sorbus. Sterile. Parmi.ua elegantula (Zahlbr.) Ras. var. americana Ras., var. nov. Thallus opacus, obscure olivaceo-fuscescens, isidiosus, K = , Cl=, K(C1)=. Isidia sub- cyliudrica, curta, conooloria. Apothecia desunt. — British Columbia: Kamloops, on a Pseudotsuga in semi-desert. Parmelia pubescens (L.) Vain. var. reticulata Cromb. British Columbia: Kamloops, on a stone in semi-desert. Sterile. Thamnolia vermicularis (Sw.) Schaer. Alaska (regio alpina), on the ground between tufts of Cornicularia divergens and Cladonia rangiferina. Sterile. SlPHULA ceratites (Wahlbg.) Fr. British Columbia: Dorreen, on stones in moist mountain forest. Sterile. Stereocaulon tomentosum Fr. British Columbia: Dorreen, on branches of T8\iga; Alberta: Lake Louise, on mossy earth; Alaska (regio alpina), between Comic- ularia divergens and Cetraria chrysantha. Rarely fertile. Lecania dimera (Nyl.) Th. Fr. British Columbia: Aleza Lake, very common on the trunks of Populus. Fertile. Squamaria alphoplaca (Wahlbg.) Dub. Thallus orbicularis, radiatus, griseus, K-+- partim rubescens. Medulla I — . Apothecia ca. 1 mm. lata, nigra, nuda vel tenuiter pruinosa. Sporae 9, 0.5 /x, 8 : nae. — U. S. A.: Cody, Wyoming, on sand- stone in a cactus desert. Aspicilia calcarea (L.) Mudd. With the previous on sandstone in cactus desert, in Cody. Fertile. Lecanora coilocarpa (Ach ) Nyl. New Brunswick: Dalhousie, on a rail fence. Fertile. Lecanora umbrina (Ehrh.) Rohl. New Brunswick: Dalhousie, on a stone in a field. Fertile. Lecanora symmicta Ach. Thallus areolato-verruculosus, mox totus sorediosus, virescens, K — , C1+ aurantiaco-rubescens. Apothecia livida, mox convexa, fere immarginata. — New Brunswick: Dalhousie, on a wood rail fence. I have found the same lichen — not in any way to be confused with Biatora sym- mictera (Nyl.) lifts. — in the summer of 1931, on a wooden fence in Petsamo, Finnland. Except for this I have not seen this central European and North American species. Lecanora subintricata Nyl. British Columbia: Kamloops, Fish Lake, on trunks and branches of Virtus Murray ana. Fertile. Ochrolechia upsaliensih (L.) Mass. Thallus K — , CI— , K(C1)— . Epi- thecium K(C1)— . Apothecia 1-2 mm. lata, planiuscula vel urceolata, pruinosa. Margo crassa, subrugosa. — British Columbia: on rotting vegetable remains in somewhat shaded positions in the semi-desert. Phlyctis argena (Ach.) Flot. New Brunswick: Dalhousie, on trunk of Abies sp. Sterile. Pertusaria pertusa (L.) Tuck. New Brunswick: Dalhousie, together with previous species on Abies. Fertile. Pertumaria xanthostoma (Smrft.) Fr. Thallus verruculoso-inaequalis, K — , K(C1) — . Apothecia 0.5 mm. lata, punetiformia, pallido-lutescentia. Excipulum K+, sanguineo-rubescens. Sporae 4: nae, ellipsoideae, incoloratae, 58-66 X 26-40 n. Hymenium 1+ caerulescens. — New Brunswick: Dalhousie, on a Thuja trunk. Pektusaria multipuncta (Turn.) Nyl. British Columbia: Aleza Lake, on the trunk of an Abies. Fertile. Pkrtifsaria faginea (L.) Vain. /\ amara (Ach.) Nyl. 1933] RASANEN — CONTRIBUTION TO LICHEN FLORA OF AMERICA 15 Thallus albidus, partim albo-sorediosus, K-f demum rubescens, K(C1)+ mox violascens. — With the previous species on an Abies at Aleza Lake. Sterile. Xanthoria polycarpa (Hoffm.) Flag. New Brunswick: Dalhousie, on a Sorbus and a rail fence. Fertile. Placodium elegans (Link) DC. New Brunswick: Dalhousie, on a field stone. Fertile. Placodium ferrugineum (Huds.) Rabenh. New Brunswick: Dalhousie, on the fibrous trunks of a Thuja. Fertile. Placodium xanthostigmoideum Ras., n. sp. Thallus totus leproso-granulosus, citrinus vel aurantiaco-citrinus, K-f violascens. Similis C andelariellae xantho- stigmae (Pers.) Lettau, sed thallus K-f. — New Brunswick: Dalhousie, on the trunks of Betula, Abies, and Thuja. Sterile. Placodium chrysodetum Vain, also has leprous thallus and positive K (violascens) reaction, but this lichen is exclusively confined to stones and may grow over dead mosses on stones. Placodium citrinum (Hoffm.) Hepp., a typical calcareous lichen, may also grow on trees and wooden houses which are impregnated with chalk dust, but is not entirely leprous and its color is " flavo-citrina vel cerina" and it frequently has apothecia. Physcia Clementiana (Ach.) Kickx. Ph. astroidea Nyl. Thallus albidus vel canus in centro granuloso-sorediosus, K+ lutescens. — New Brunswick : Dalhousie, on the bark and on the peeled trunks of Picea. Sterile. Physcia stellaris (L.) Nyl. New Brunswick: Dalhousie, with the previous species on Picea. Sterile. Physcia stellaris var. rosulata (Ach.) Nyl. New Brunswick: Dalhousie, on a trunk of Betula. Fertile. Physcia caesia (Hoffm.) Nyl. New Brunswick: Dalhousie, with Placodium elegans on a field stone. Sterile. Physcia muscigena (Ach.) Nyl. British Columbia: Kamloops, on the ground between mosses and grasses in a semi-desert. Sterile. Physcia muscigena f. lenta (Ach.) Vain. British Columbia: Kamloops, with the previous species on the ground in semi-desert. Sterile. Physcia muscigena f . squarrosa (Ach.) Lynge. Thallus 6-8 cm. latus, imbricatus, albo-pruinosus; laciniae curtae, obtusae, adscendentes. Apothecia 1-3 mm. lata, plana, albo-pruinosa, marginata. Sporae 15-26 X 8-15 /x, fuscae. — British Colum- bia: Kamloops, with the previous species on the ground in semi-desert. Physcia grisea (Lamy) Zahlbr. var. semifarrea (Vain.) Lynge. British Co- lumbia: Kamloops, on the ground between grasses and Selaginella. Sterile. Physcia pyrithrocardia (Mull.-Arg.) Ras. Ph. adglutinata var. pyrithrocardia Mull.-Arg. Thallus centrum versus tote isidioso-sorediosus, viridi-griseus, intus erythrinus, K = . — New Brunswick: Dalhousie, on the trunk of Picea. Sterile. Physcia endophoenicea (Harm.) Ras. Sicut Ph. obscura (Ehrh.) Nyl. f. ciliata (Hoffm.) Lynge, sed thallus fusco-niger et intus pulchre rubescens. Thallus KqF violascens. — New Brunswick : Dalhousie, on trunk of Betula, with Placodium xantho- stigmoideum. Sterile. Rinodina demissa (Flk.) Mass. New Brunswick: Dalhousie, on a field stone. Fertile. Buellia disciformis (Fr.) Mudd var. minor (Fr.) Ras. New Brunswick: Dal- housie, on a Populus trunk. Fertile. [Vol. 20 10 ANNALS OF THE MISSOURI BOTANICAL GARDEN Uuklma disciforms var. iNSiGNis (Naeg.) Nyl. f. muscorum (Schaer.) Ras. British ( 'olumbia: Kamloops, on mosses growing on stones in a semi-desert. Fertile. Bubllia major (DN.) Mass. New Brunswick: Dalhousie, on trunks of Sorbus, Picea, and Abies. Fertile. It helm A punctiformin (lloffm.) Mass. f. chlohopolia (Fr.) Vain. British Columbia: Kamloops, on the trunk (bark) of a very well-lighted Pseudotsuga. Fertile, I'eltighra variolosa (Mass.) Korb. Subsimilis Pcltigerae aphthosae sed thallus subtus distincte nervosus et superne marginem versus subpruinosus. — British Columbia: Field, on the ground between mosses and grasses. Sterile. Peltigera venosa (L.) Korb. British Columbia: Dorreen, on mineral soil in shaded forest. Fertile. In the dozen individuals from Dorreen the apothecia are much smaller (about 1 mm.) than in the Finnish specimens, so that one might regard the American species as a new variety (var. microcarpa Ras.). PsLTiOEBA horizontals (L.) Hoffm. British Columbia: Prince George, Six Mile Lake, on mossy stone walls. Fertile. Peltigera maurizii Gyeln. (Hedwigia 68: 1. 1928). Thallus circiter 6 cm. latus, superne laevis, nitidus, epruinosus et etomentosus, cinereus vel cinereo-fusces- c(mis, subtus malaceaeforme nervosus, obscuro-fuscus, marginem versus pallidior; rhizinae nigro-fuscae, fascieulatae, obsoletae. Apothecia horizontalia, plana vel concava, badio-fusca, 3-5 mm. lata; margo subintegra. Sporae 26-43 X 3.5-6 n, fusiform es, 3-septatae, hyalinae. Hymenium 80-83 m crassum, sordide-hyalinum, 1+ caerulescens. — British Columbia: Field, on ground between mosses and other ground lichens. This easily recognizable lichen is a form intermediate between Peltigera malacea and P. horizontalia. Peltigera malacea (Ach.) Fr. British Columbia: Valemount, on an Arctosta- phylos and lichen heath. On the ground. Sterile. Peltigera canina (L.) Willd. British Columbia: Kamloops, on mossy stones in semi-desert. Fertile. Peltigera canina var. membranacea Ach. British Columbia: Field, with Peltigera Maurizii and P. variolosa, on the ground in a forest of Picea. Fertile. Peltigera canina var. nuomensis (Gyeln.) Rite. /\ ituomensis Gyeln. (Magyar Bot. Lap. 29: 34. 1930). British Columbia: Blue River, on the ground in a burnt-over Myrtillus forest. Fertile. Peltigera rufescens (Weis) Humb. British Columbia: Kamloops, on the ground in semi-desert. Fertile. Pkltigera microphylla (Anders.) Gyeln. P. perfida Gyeln. Thallus superne cinereo-fuscescens, laevigatus, nitidus, subtus polydactylaeforme venosus, marginem versus squamuloso-isidiosus. Apothecia junior involuta. — New Brunswick: Dalhousie, on ground in coniferous forest. Peltigera lepidophora (Nyl.) Bitt. Alberta: Wainwright, on calcareous earth, together with Cladonia cariosa, in a desert. Rare and sterile. Peltigera Nylanderi Gyeln. (Bot. Kozlem. 24: 137. 1927). Thallus pusillus, 12 cm. latus, firmus, monophyllus, laciniatus, ad marginem bene limbiformiter sored iosus, subtus caninaeforme venosus. — British Columbia: Kamloops, between mosses on stones in semi-desert. Sterile. Nephromium subparile Gyeln. (Magyar Bot. Lap. 29: 24. 1930). Thallus 1933] RASANEN — CONTRIBUTION TO LICHEN FLORA OF AMERICA 17 superne et ad marginem bene granuloso-sorediosus, etomentosus (glaber), subtus glaber, K = . — British Columbia: Kamloops, on stones in semi-desert. Sterile. Nephromium laevigatum Ach. var. papyraceum (Hoffm.) Nyl. Thallus pusillus, tenuis, superne glaucus, K = , subtus albido-pallens, glaber. Apothecia majora, 3-8 mm. lata, horizontalia, fusca; margo subintegra. Sporae pallidae, 16-21 X 4-5 /i, 1- vel 3-septatae, oblongo-fusiformes. — British Columbia: Aleza Lake, on branches of Picea in the forest. Nephromium canadense Ras., n. sp. Thallus 5 cm. latus, subrigidus, coriaceus, bene laciniatus, ad marginem dentatus vel lacunate dentatus, superne sublaevigatus, apicem versus scabroso-areolatus, passim leviter hirsutus vel totus nudus, griseo- pallidus vel glauco-fuscescens, subtus tomentellus, pallido-fuscescens. Apothecia vulgaria, 3-6 mm. lata, convoluta, fusca; margo dentata. Sporae 3-septatae, pallidae, oblongae vel oblonge fusiformes, 14.5-21 X 7-8 m- Hymenium 70 m crassum. 1 + caerulescens. Gonidia nostocoidea. — British Columbia: Aleza Lake, together with Nephromium laevigatum var. papyraceum and Cyanisticta Hookeri var. septentrionalis , on branches of Picea in forest. Solorina saccata (L.) Ach. British Columbia: Field, on a tree base. Fertile. Lobarina verrucosa (Huds.) Gyeln. Sticta scrobiculata (Scop.) Ach. New Brunswick: Dalhousie, on a tree trunk, rare. Sterile. Lobaria pulmonaria (L.) Hoffm. Thallus ad marginem sorediosus, non isidiosus; medulla et soredia K+ flavescentia. — New Brunswick: Dalhousie, on Abies and Betula trunks. Sterile; British Columbia: Aleza Lake, on branches and trunks of Picea. Fertile. Lobaria linita (Ach.) Rabenh. Thallus esorediatus, K = . — Alaska (regio alpina), on the ground between Cladonia species and mosses. Sterile and rare. Lobaria oregana (Tuck.) Miill.-Arg. Thallus majus, 30 cm. longus et 3-10 cm. latus, dichotome laciniatus, lacunoso-reticulatus, viridi-glaucus, esorediatus, basim versus demum squamulosus; laciniae in apicibus rotundata. Apothecia vulgaria, in centro aut submarginalia, plana aut demum convexa, castaneo- vel rufo-fusca; margo tenuis. Sporae 40-61 X 6-8 n, hyalinae, fusiformes, 1-septatae. Hymenium 100-120 m crassum, incoloratum, I +, caerulescens. Hypothecium 60 m crassum, K + lutescens. Thallus K=F lutescens, CI = , K(C1) =, I = . — British Columbia: Hazleton, on Pinus Murrayana, Dorreen, on Pseudotsuga, and Vancouver Is., Lake Cowichan, on trunks of Thuja and Tsuga. Stictina fuliginosa (Dicks.) Nyl. British Columbia: Aleza Lake, common on branches of Picea in forest. Sterile. Cyanisticta Hookeri (Bab.) Ras. var. septentrionalis Ras., var. nov. Thallus superne glauco-cinereus, subnitidus, scrobiculatus, isidiosus, subtus obscure rhizino- sus. Pseudocyphellia minora, papilliformia, albida. Gonidia nostocoidea. — British Columbia: Aleza Lake, on branches of Picea. Sterile. Collema nigrescens (Huds.) Ach. New Brunswick: Dalhousie, on Populus trunks. Sterile. Leptogium tremelloides (L.) Fr. New Brunswick: Dalhousie, on Abies trunks. Sterile. Leptogium saturninum (Dicks.) Mass. British Columbia: Kamloops, on side of a stone in semi-desert. Sterile. Pilophoron aciculare (Tuck.) Nyl. Thallus 2.5-3.5 cm. longus, simplex vel parce ramosus, corticatus, viridis, K+ lutescens. Apothecia sphaeroidea vel sub- [Vol. 20 18 ANNALS OP THE MISSOURI BOTANICAL GARDEN conica, vulgaria, nigra vel caeruleo-nigra. Sporae hyalinae, oblongo-fusiformes, simplices, 18-21 X 5-6 m- Ei)ithecium caeruleum, K+ smaragdulum. Hypothe- cium fuligineum. Hymeriium 60-130 m, crassum, incoloratum, 1+ caerulescens.— British Columbia: Dorreen, frequent on stones between plant remains (fallen needles, etc.) in mountain forest. Cladonia rangiferina (L.) Rabenh. Alaska (regio alpina), on ground. Sterile. Cladonia rangiferina f. nivea Ras. Thallus niveus, K+ lutescens. — British Columbia: Dorreen, on an arid heath in Pinus Murrayana forest. Sterile. Cladonia kilvatica (L.) Rabenh. Alaska (regio alpina), on ground; British Columbia: Dorreen, on an arid heath among Pinus Murrayana. Fertile. Cladonia silvatica var. mitis (Sandst.) Ras. British Columbia: Valemount, on an arid heath with Arctostaphylos Uva-ursi. Sterile. Cladonia alpestris (L.) Rabenh. Alberta: Lake Louise, on an Empetrum heath. Common. Sterile. Cladonia bacillaris Nyl. New Brunswick: Dalhousie, on a very rotten rail fence. Fertile. Cladonia cristatella Tuck. New Brunswick: Dalhousie, in same situation as previous species. Fertile. Cladonia deformis Hoffm. Alberta: Lake Louise, on an arid heath between other Cladonia species and mosses. Sterile. Cladonia uncialih (L.) Fr. Alaska (regio alpina), on ground; British Columbia: Valemount, on an Arctostaphylos-Cladina heath. Fertile. Cladonia crispata (Ach.) Flot. var. infundibulifera (Schaer.) Vain. British Columbia: Valemount, with previous species on an Arctostaphylos-Cladina heath. Fertile. Cladonia crispata var. elegans (Del.) Vain. British Columbia: Blue River, on a burnt-over V actinium Myrtillus heath. Fertile. Cladonia cenotea (Ach.) Schaer. var. crossota (Ach.) Nyl. British Columbia: Valemount, on an Arctostaphylos-Cladina heath. Sterile. Cladonia oariosa (Ach.) Spreng. f. cribrosa (Wallr.) Vain. Alberta: Wain- wright, on ground in desert with Peltigera lejridophora. Fertile. As both species grow on calcareous soil, one may conclude that the soil in these deserts is calcareous. Cladonia gracilis (L.) Willd. var. dilatata (Hoffm.) Vain. New Brunswick: Dalhousie, on a very rotten rail fence; British Columbia: Golden, on a sand heath; Valemount, on an Arctostaphylos-Cladina heath; Alberta, Lake Louise, on an arid heath. Fertile. Cladonia gracilis f. anthocephala Flk. British Columbia: Field (regio sub- alpina), on a V actinium Myrtillus heath; Blue River, on a V. Myrtillus heath, which had been burnt over. Fertile. Cladonia elongata (Jacq.) Hoffm. Alaska (regio alpina), on mossy ground. Sterile. Cladonia elongata var. ecmocyna (Ach.) Ras. Podetia cornuta vel subcornuta, saepe dilatata, scyphifera, glauco-cinerea, K — pulchre lutescentia.- — Alberta: Lake Louise, on a dry heath; British Columbia: Field (regio alpina), on heath under Phyllodoce, etc. Fertile. Cladonia cornuta (L.) Schaer. New Bruiuswick: Dalhousie, on rotten stump of an Abies. Sterile. Cladonia verticillata Hoffm. New Brunswick: Dalhousie, on ground. Sterile. Cladonia degenerans (Flk.) Spreng. British Columbia: Valemount, on an Arctostaphylos heath. Fertile. 1933] RASANEN CONTRIBUTION TO LICHEN FLORA OF AMERICA 19 Cladonia subcervicornis (Vain.) DR. f. turgescens Magn. (Lich. Sel. Scand. Exs. II, No. 34. 1929). British Columbia: Kamloops, on a desert. Fertile. Cladonia pyxidata (L.) Fr. var. neglecta (Flk.) Mass. British Columbia: Kamloops, on edge of desert. Sterile, Cladonia chlorophaea (Flk.) Spreng. British Columbia: Kamloops, on ground in desert; Alberta: Wainwright, in desert. Sterile. Cladonia fimbriata (L.) Fr. var. simplex (Weis) Vain. f. minor (Hag.) Vain. New Brunswick: Dalhousie, at base of Picea; British Columbia: Kamloops, in desert. Sterile. Cladonia fimbriata var. simplex f. major (Hag.) Vain. British Columbia: Aleza Lake, on Picea base. Sterile. Cladonia fimbriata var. apolepta (Ach.) Vain. New Brunswick: Dalhousie, on a very rotten rail fence. Fertile. Bacidia minuscula Anzi var. Beckhausii (Korb.) Vain. Thallus obsoletus. Apothecia minutissima, 0.2-0.3 mm. lata, globosa, tuberculata, nigra, tenuiter pruinosa. Sporae juvenes, 12-18 X 2 ju, 1- to 3-septatae, bacillariae, incoloratae. Hymenium obscure olivaceum, 1+ caerulescens, deinde sordide rubescens, 40 m crassum. Epithecium K+ violascens. Hypothecium pallidum. — British Columbia: Aleza Lake, on bark of Abies with Biatora helvola and Opegrapha sp. Bacidia fuscorubella (Hoffm.) Am. British Columbia: Aleza Lake, on trunk of a Picea. Fertile. Psora decipiens (Ehrh.) Korb. British Columbia: Field (regio subalpina), on sandy ground. Fertile. Catillaria tricolor (With.) Th. Fr. British Columbia: Aleza Lake, occasional on twigs of Pinus Murrayana. Fertile. Biatora symmictera (Nyl.) Ras. New Brunswick: Dalhousie, on trunks and branches of Thuja, Abies, and Sorbus. Fertile. Biatora helvola (Korb.) Th. Fr. British Columbia: Aleza Lake, on bark of Abies. Fertile. Lecidea goniophila Flk. U. S. A.: Cody, Wyoming, on sandstone in light in a cactus desert. Fertile. Lecidea lapicida (Ach.) Vain. var. seriata Th. Fr. U. S. A.: Cody, Wyoming, in same situation as previous species. Fertile. Mycoblastus melinus (Krmplhbr.) Hellb. Thallus albido-cinereus, verruculosus, esorediatus, K+ lutescens. Apothecia nigra, opaca, 0.5-1.5 mm. lata, convexa, immarginata. Sporae binae, hyalinae, ellipsoideae, 46-55 X 25-32 /x. Hypothecium pallidum. — British Columbia: Aleza Lake, on barkless, dry, decorticated twigs of Picea. Mycoblastus sanguinarius (L.) Norm. Thallus viridi-albidus, K— , intus sanguineus. Apothecia nigra, nitida. Sporae 46-61 X 25-31 m, solitariae. Hypo- thecium crasse sanguineum. — British Columbia: Aleza Lake, on Betula and Abies trunks. Fertile. Mycoblastus sanguinarius var. Dodgeanus 1 Ras., var. nov. Thallus albidus, verruculoso-inaequalis, esorediatus, tenuis vel mediocris, K+ auratus, CI— , I — , intus albidus. Apothecia 1 mm. lata, atra, convexa, opaca, immarginata. Hypo- thecium tenuiter sanguineum vel pallido-flavescens. Sporae solitariae, magnae, oblongo-ellipsoideae, 85-92 X 26-36 \i. — British Columbia: Aleza Lake, common on bark of Abies lasiocarpa; Prince George and Six Mile Lake, on Betula. 1 In honor of Dr. C. W. Dodge. [Vol. 20 20 ANNALS OF THE MISSOURI BOTANICAL GARDEN ACABOSPORA Schleicheri (Ach.) Mass. British Columbia: Kamloops, on sandy ground in a desert. Fertile. Acarohpoka molybdina (Wnhg.) Trevis. U. S. A.: Cody, Wyoming, on sand- stone in cactus desert. Fertile. Acabohpoka peltasticta Zahlbr. U. B. A. I Cody, Wyoming, with previous species on exposed sandstone in cactus desert. Fertile. Diplohchintes bryophiloideh (Nyl.) Zahlbr. Alberta: Wainwright, on ground over mosses and plant remains. Fertile. DiPLOsmiHTEH canadensis Riis., n. sp. Thallus griseo-cinereus, areolata-verru- culosus, mediocris, Kl K(C1) haud roseus, I *. Apothecia 0.5-1 mm. lata, urceolata, atra, haud pruinosa, marginata; margo erenata. Sporae 4 : nae, fuscae vel demum nigrae, muriformes, 19.5-27.5 X 11-13 p. Ilymenium 1+ fulvescens. Sicut D. scruposus var. terrestris sed thallus cum KOI I rubescens. — British Columbia: Kamloops, on sandy ground in desert. G ha phis scripta (L.) Ach. New Brunswick: Dalhousie, on Abies trunks; British Columbia: Aleza Lake, on Abies lasiocarpa. Fertile. Sphaekophorui globosus (Huds.) Vain. var. lacunosus Tuck. Thallus 7 cm. altus, bene ramosus, pallido- vel olivaceo-fuscus, K = , Cl=, K(C1)=, I=F distincte caerulescens. Rami lacunoso-foveolati. Apothecia globosa, subterminalia. — British Columbia: Hazleton, on ground. Fertile. Sfhaeroimiohus Tuckermanii Riis., n. sp. Thallus 5-8 cm. altus, parce ramosus, albidus vel olivaceo-albidus, passim albido-roseus, K — , CI — , K(C1) — , It dilute caerulescens, cum ramis primariis, cylindricis, 0.5-1 mm. latis, laevibus et cum ramulis adventiciis, 1-5 mm. longis, sat tenuibus et parcis. Apothecia vulgaria, terminalia, globosa, 1-2 mm. lata. Massa sporalia, nigra. Sporae 8 : nae, sphaer- oideae, 5-8 m, viridi-atrae vel fusco-nigrae. Asci anguste cylindrici. — British Co- lumbia: Hazleton, with previous species on ground between mosses and plant re- mains. Staurothele circinata Tuck. Thallus fusco-niger, areolato-diffractus, ad marginem subradiatus. Apothecia concoloria. Sporae fuscae, binae, murales, oblongo-ellipsoideae, 39-52 X 21-25 m- Gonidia hyrnenialia, oblongia, viridia, 8-9.2 X 4 p. — U. S. A.: Cody, Wyoming, on sandstone with Acarospora molybdina and Squamaria alphoplaca in a cactus desert. •« 1933] • 4 • « RASANEN — CONTRIBUTION TO LICHEN FLORA OF AMERICA 21 Genus Index to Genera Page Genus Page Acarospora 20 Nephromopsis 13 Alectoria 10 Ochrolechia 14 Aspicilia 14 Parmelia 13 Bacidia 19 Peltigera 16 Biatora 19 Pertusaria 14 Buellia 15 Phlyctis 14 Catillaria 19 Physcia 15 Cetraria 11 Pilophoron 17 Cladonia 18 Placodium 15 Collema 17 Psora 19 Cornicularia 11 Ramalina 11 Cyanisticta 17 Rinodina 15 Diploschistes 20 Siphula 14 Graphis 20 Solorina 17 Lecania 14 Sphaerophorus 20 Lecanora 14 Squamaria 14 Lecidea 19 Staurothele 20 Leptogium 17 Stereocaulon 14 Letharia 11 Stictina 17 Lobaria 17 Thamnolia 14 Lobarina 17 Usnea 9 Mycoblastus 19 Xanthoria 15 Nephromium 16 S lm ASTRANTHIUM AND RELATED GENERA ESTHER LOUISE LARSEN Formerly Assistant Professor of Botany, University of Montana Formerly Jessie R. Barr Fellow in Botany, Washington University Monographic studies of Aphanostephus and Achaetogeron have made it necessary to investigate some of the related genera, particularly as to the value of certain morphological characters which are used in generic differentiation. The genera are quite similar in habit but the pappus, which furnishes diagnostic characters, varies greatly within the group and is on the whole inconspicuous. For this reason it is necessary to make microscopic studies of the minute achenial and pappus characters upon which the differentiation of genera is primarily dependent. In this connection it was desirable to study Astranthium and Keerlia in detail; and it seems advisable to put on record the results which have been obtained relative to these genera. Astranthium, a genus of the Compositae belonging to the tribe Astereae, was described by Nuttall in 1841. * It contained but one species, A. integrif olium , based on Bellis integrifolia Michaux. 2 The 'Flora of North America,' published by Torrey and Gray in 1842, relegated Astranthium to synonymy. Since that time American and Mexican species have been merged with the genus Bellis 3 which is indigenous to the Old World and especially to Europe. Three South American species have been described by Vellozo 4 as occurring in Brazil. Two of these, Bellis campestris Veil, and B. pedunculata Veil., may well be members of the genus Spilanthes. The relationship oi Bellis scandens Veil., is unknown to me. third and for it has been cultivated in the north Atlantic states where it has become naturalized. The American species which have been referred hitherto to the genus Bellis are so strikingly different that it seems strange the two generic elements should have been regarded 1 Nutt. Trans. Am. Phil. Soc. N. S. 7: 312. 1841. 2 Michx. Fl. Bor. Am. 2: 131. 1803. 3 Torr. & Gray, PL N. Am. 2: 189. 1842. * Veil. Fl. Flum. 8: pi. 124-126. 1827; text, pp. 338, 359, ed. 1881. Issued April 29, 1933. Ann. Mo. Bot. Gard., Vol. 20, 1933. (23) [Vol. 20 24 ANNALS OF THE MISSOURI BOTANICAL GARDEN for such a long time as congeneric. The range of Bellis in North America is far to the north of Astranthium integrif olium (Michx.) Nutt., whose northernmost limit of distribution is in Kentucky. Bellis perennis L., on the other hand, is reported from the islands of St. Pierre and Miquelon, off the coast of Newfoundland, south to Pennsylvania, and in northwestern America north to Vancouver Island. Because of the difference in morphological characters it seems advisable to recognize Astranthium as a distinct genus. Six Mexican species have been described under Bellis, all of which are here transferred to Astranthium, except B. Garciae Blake. The first Mexican species was described by Gray in 1S52 as Bellis mexicana,* and in 1881 he transferred Brachycome xanthocomoides* to that genus. Since then four additional species have been published, namely, B. purpurascens Rob., 7 B. ortho- poda Hob. & Fern., 8 B. mima, and B. Garciae <>f Blake. 9 Another species is added to this group by the transfer of Keerlia mexicana Gray ex Wats. 10 to Astranthium where it receives the new name Astranthium xylopodum Larsen, because of the earlier Bellis mexicana Gray. Only one species, Astranthium integrif olium, occurs in the United States. In 1836 Rafinesque described three species, Bellis parvifolia, Bellis nutans, and Bellis ciliata. 11 The first two are considered synonymous with Astranthium integrif olium and the last is regarded as a variety of that species. The genus Keerlia, which is closely related to Astranthium, was first described by De Candolle 12 in 1 836. It contained three species, K. linearif olia , K. ramosa, and K. skirrhobasis . All three species have been transferred to other genera since that time. Keerlia linearif olia, which would ordinarily be interpreted as the generic type, was based on two distinct plants, one collected by Alaman and the other by Schiede. The former was referred to the genus Xanthocephalum by Dr. J. M. Greenman who pub- 6 Gr.iv, Smiths. Contr. [PL Wright, |>t. 1| 3: 93. 1852. •Gray in Hemsl. Biol. Cent.-Am. Bot. 2: 118. 1881. 7 Rob. Proc. Am. Acad. 27: 172. IK!) 8 Rob. A I'Vrn. Pro< Am. Acad. 30: 117. 1894. • Blake, Contr. U. S. Nat. Herb. 22: 593. 1924. 10 Gray ex Wats. Proc. Am. Acad. 22: 422. 1887. 11 Raf. New Fl. Am. 2:23. 1836. '•-' D. C. Prodr. 5: 309. 1830. 1933] LARSEN — ASTRANTHIUM AND RELATED GENERA 25 lished the new combination X anthocephalum linearif olium (DC). Greenm. 13 The latter, namely, the Schiede plant, on which Brachycome xanthocomoides was based and cited in synonymy by De Candolle, was transferred by Hemsley to the genus Bellis and published as Bellis xanthocomoides Gray ex Hemsley. 14 Keerlia ramosa 16 and Keerlia skirrhobasis 16 have both been transferred by Gray to the genus Aphanostephus. Since the original publication of the genus by De Candolle the following species have been described: Keerlia bellidifolia Gray & Engelm., 17 Keerlia effusa Gray, 18 and Keerlia mexicana Gray ex Wats. 19 Of these three species only the first two are retained in Keerlia, the third is transferred to Astranthium. Although all of the species originally described by De Candolle in Keerlia have been excluded from that genus, nevertheless it seems advisable to use Keerlia as the generic name for the residue, namely, for those species which have been described subsequently and still remain in that genus. The historical name is thus retained; and the genus is delimited in accordance with Gray's treatment in the ' Synoptical Flora of North America'. 20 More- over, the name Bourdonia, proposed by Greene, 21 falls to syn- onymy General Morphology The American genus Astranthium is morphologically distinct from Bellis of the Old World with which it was merged in 1842. In habit the genera are quite different. Bellis produces long naked monocephalous peduncles from rosulate clusters of basal leaves, whereas the stems of Astranthium are usually branched and seldom entirely naked. The involucre of Astranthium is composed of two or three series of bracts which are usually membranaceous margined. 13 Greenm. Field Mus. Nat. Hist. Publ. Bot. Ser. 2: 345. 1912. 14 Gray in Hemsl. Biol. Cent.-Am. Bot. 2: 118. 1881. 15 Gray, Proc. Am. Acad. 16: 81. 1881. 16 Gray, Smiths. Contr. [PI. Wright, pt. 1] 3: 93. 1852. 17 Gray & Engelm. Proc. Am. Acad. 1 : 47. 1848. 18 Gray, Bost. Jour. Nat. Hist. [PI. Lindh. pt. 2] 6: 222. 1850. 19 Gray, Proc. Am. Acad. 22 : 422. 1887. 20 Gray, Syn. Fl. N. Am. I 2 : 164. ed. 2, 1886 and 1888. 11 Greene, Erythea 1 : 207. 1893. 26 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 20 The involucral bracts of Bellis differ in that they are subuniseriate, more or less united at the base, and distinctly foliaceous through- out. On the whole they are broader than those of Astranthium and less acute. A marked contrast between the two genera is to be found in the receptacle. In Astranthium it is a low, convex, cushion-like structure with alveolate surface of a light creamy or whitish color. The receptacle of Bellis is distinctly conical. It has a smooth, almost shiny, dark greenish-brown surface which is broken only by the white spots indicating points of achenial attachment, and by depressions which are due to shriveling of the subepidermal tissue (pi. 2, figs. 1 and 7). The corollas of the disk- and ray-flowers are very similar in these genera. The ray-flowers of Astranthium may be said to be proportionately longer than those of Bellis. The stamens in both cases are typically asteroid (pi. 2, figs. 2, 3, 4, 8, 9, 10). The achenes of the genera are similar in outline and size. Bellishiis achenes with distinct margins, while t hose of A str anthium are without distinct margins and are usually more or less pubes- cent with glochidiate-tipped hairs. The pistils are similar in size but differ in the characters of their style-branches. Astranthium has long, slender, acute style- branches in contrast to those of Bellis, which are only about one- half as long as broad, thick, and obtuse (pi. 2, figs. 5, 6, 11, 12). The pappus is entirely lacking in Bellis and this is also the most usual condition in Astranthium. There are, however, certain exceptions. Astranthium xanthocomoides has a slight indication of a ring-like crown. The ring-like crown is more noticeable in A. orthopodum and becomes conspicuous in A. mexicanum var. chihuahuense where it is somewhat fluted. This vestigial pappus when present forms an unbroken ring-like crown which is entirely without lacerate margins commonly found in related genera. < Jeographical Distribution The only representative of the genus Astranthium occurring in the United States is Astranthium integrifolium with its two varieties ciliatum and rosulatum. The species extends from 1933] LARSEN — ASTRANTHIUM AND RELATED GENERA 27 •2 3 CO a o 3 .2 bC O o a faC 3* si a 8 8-S to °° °3 o3 H d (1) 3 o O 00 ,g o 28 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 20 Georgia and Kentucky to western Oklahoma and Kansas. The varieties occur in Oklahoma, Arkansas, and Texas. Mexican representatives of Astranthium are scattered through- out Mexico from Chihuahua to Chiapas. There is very little material in American herbaria representing Mexican species. Astranthium mimum is known only from the locality in which it was collected originally. Astranthium purpurascens was found by C. G. Cringle in San Luis Potosi, but the same species was also collected previously by Dr. A. Ghiesbreght in Chiapas. Astran- thium xylopodum is reported only from eastern Jalisco where it has been collected by both Dr. Edward Palmer and C. G. Pringle. Astranthium orthopodum occurs in the mountains of Chihuahua and in Durango. The oldest Mexican species and the one which is best represented in herbaria is A. mexicanum. It extends from ]Iidalgo to central Oaxaca; and a variety, chihuahuense , occurs in northwestern Chihuahua. Astranthium xanthocomoides has been reported from Hidalgo and Vera Cruz. The scattered distribution of Astranthium in Mexico is prob- ably an indication of the incomplete representation of the flora of that country in herbaria. Further collections will undoubtedly extend the distribution areas and also add to the number of species. Two species of the genus Keerlia are recognized and both of these occur in south-central Texas. It is not at all unlikely that they will be found in adjacent Mexico. The distribution of the various species representing Astranthium and Keerlia is shown on the accompanying map. Acknowledgments The writer wishes to express her appreciation to Dr. George T. Moore, Director of the Missouri Botanical Garden, for the use of the library and herbarium of that institution, and especially to Dr. J. M. Greenman for advice and assistance. Dr. B. L. Robinson and Dr. W. R. Maxon kindly loaned material which was necessary to the study of these genera. Abbreviations The specimens cited in this paper are deposited in the herbaria which are indicated by the following abbreviations: 1933] LARSEN — ASTRANTHIUM AND RELATED GENERA 29 G = Gray Herbarium of Harvard University M = Missouri Botanical Garden Herbarium. US = United States National Herbarium. Key to the Genera A. Heads small, few-flowered Keerlia AA. Heads medium-sized, many-flowered. a. Achenes strongly compressed, not 4-angled. b. Heads subglobose; rays relatively short Egletes bb. Heads not subglobose; rays relatively long. c. Pappus absent or of a ring-like crown; ray-flowers uniseriate. d. Receptacle smooth, conical; involucral bracts sub-uniseriate . . . Bellis dd. Receptacle alveolate, convex; involucral bracts bi-tri-seriate. Astranthium cc. Pappus of a short unequally lacerated crown; ray-flowers in 2 or more series Achaetogeron aa. Achenes not compressed, subterete, or distinctly 4-5-angled. . . . Aphanostephw Key to Species of Astranthium A. Stems erect or suberect from a distinctly ligneous base. a. Leaves glabrous LA. mimum aa. Leaves more or less pubescent. b. Stems usually unbranched above. c. Leaves oblong-elliptic 8. A. xylopodum cc. Leaves oblanceolate to linear S. A. orthopodum bb. Stems usually branched above. c. Leaves dentate 4- A. mexicanum cc. Leaves entire J^a. Var. chihuahuense AA. Stems erect or procumbent from an annual or slightly ligneous base. a. Plants with long slender runners 6. A. xanthocomoides aa. Plants without long slender runners. b. Achenes glabrous 6. A. purpurascens bb. Achenes pubescent with glochidiate-tipped hairs. c. Leaves not rosulate. d. Plants not conspicuously branched 7. A. inlegrifolium dd. Plants conspicuously branched 7a. Var. ciliatum cc. Leaves rosulate 76. Var. rosulatum Taxonomy astranthium Astranthium Nutt. Trans. Am. Phil. Soc. N. S. 7: 312. 1841; Benth. & Hook. Gen. PL 2: 265. 1873, as to synonymy. Bellis Michx. and Am. authors, Fl. Bor. Am. 2: 131. 1803, not of Linnaeus; Hook. Bot. Mag. 52: pi. 8455. 1835; DC. Prodr. 5: 304. 1836; Raf. New Fl. Am. 2: 24. 1836; Torr. & [Vol. 20 30 ANNALS OF THE MISSOURI BOTANICAL GARDEN Gray, Fl. N. Am. 2: 189. 1842; Gray in Hemsl. Biol. Cent.-Am. Bot. 2: 118. 1881; Gray, Syn. Fl. N. Am. I 2 : 103. 1884, and ed. 2, 163. 1886 and 1888; Britton & Brown, 111. Fl. 3: 350. 1898, and ed. 2, 3: 402. 1913; Britton, Manual, 943. 1901, id ed. 2, 1905; Rob. & Fern, in Gray, Manual, ed. 7, 799. 1908; nail, Fl. Southeastern U. S. 1202. 1903, and ed. 2, 1913. Herbaceous, caulescent, glabrate, or pubescent annuals or perennials. petioled, linear-lanceo- late to < )bovate-spathulate, entire or dentate. Involucre 2-3- seriate, imbricated, appressed, lanceolate bracts usually with lacerately ciliate, membranaceous margins. Heads few- to many- flowered. Ray-flowers pistillate, rays 2-3-dentate. Disk-flowers tubular, perfect, corolla 5-lobed. Style-branches linear-lanceo- late, acute, papillose at the tip, stigmatic surfaces confined to the lowermost margins of the style-branches and extending one third of their length. Pappus entirely lacking or an inconspicu- ous ring-like crown. Achenes obovate, compressed, narrowed at the base, pubescent with glochidiate or emarginate hairs, rarely glabrous. Type species: Astranthium intcgrijolium (Michx.) Nutt. Trans. Am. Phil. Soc. N. S. 7: 312. 1841, which was based on Bellis integrifolia Michx. Fl. Bor. Am. 2: 131. 1803, "ad ripas rivu- lorum et in collibus umbrosis Tennassee." 1. Astranthium minium (Blake) Larsen, n. comb. Hdlis mint a Blake, Contr. U. S. Nat. Herb. 22: 594. 1924. Herbaceous perennial, 38-50 cm. high; stems several, simple, erect, monocephalous, greenish, striate, glabrous or sparsely pubescent with spreading hairs; basal leaves few, obovate- oblong, apiculate, narrowed at the base into a slender petiole, 15-20 cm. long, 2-3 cm. wide, somewhat membranous in the dried state; stem-leaves linear-lanceolate, hirsute-ciliate, lower 5-6 cm. long, 4-0 mm. wide, gradually reduced above, the uppermost 1 cm. long, 1 mm. wide; peduncles terminal, mono- cephalous, enlarged just below the head; involucre 2-2.5 cm. in diameter; bracts 2-seriate, equal, 7 mm. high, linear, acute, sparsely pilose, ciliate; ray-flowers about 00, white, fertile, 3- dentate, about 12 mm. long; disk-flowers yellow, fertile, pappus 1933] LARSEN — ASTRANTHIUM AND RELATED GENERA 31 2440-2745 none; achenes of ray and disk similar, oblong, compressed glabrous. Distribution: State of Durango, Mexico. Known only from type specimen. Specimens examined: Durango: from Sierra Madre, 30 miles north of Guanacevi, alt. Aug. 18, 1898, Nelson 4786 (US, No. 332836, type). 2. Astranthium xylopodum Larsen, n. name. Keerlia mexicana Gray, Proc. Am. Acad. 22: 422. 1887, not Bellis mexicana Gray, Smiths. Contr. [PI. Wright, pt. 1] 3: 93. 1852. Perennial from a woody base ; stems several, simple, or slightly branched near the top, striate, hirsute with spreading pubescence ; peduncles 10-14 cm. long, naked or bearing 1-2 small linear- lanceolate bracts near the inflorescence, hirsute-pubescent; leaves oblong-elliptic, evenly pubescent with appressed hirsute hairs, margins strongly ciliate; involucre 1-2 cm. in diameter; bracts linear-lanceolate, pubescent, margins lacerately ciliate, somewhat membranaceous; ray-flowers fertile, about 20; pappus lacking; achenes 4-nerved, glabrous. Distribution: known only from the State of Jalisco, Mexico. Specimens examined: Rio Blanco, July 1886. Edward Palmer Pringl 3. Astranthium orthopodum (Rob. & Fern.) Larsen, n. comb. Bellis orthopoda Rob. & Fern. Proc. Am. Acad. 30: 117. 1894. Perennial from a short, thick rootstalk; stems several, de- cumbent or suberect, 10-20 cm. high, simple or somewhat branched, appressed-pubescent, monocephalous ; leaves thick, entire, appressed-pubescent, basal leaves oblong-spathulate, 3-4 cm. long, 3-5 mm. broad, stem-leaves linear-lanceolate, erect, gradually reduced toward the inflorescence; peduncles 3^1.5 cm. long, appressed-pubescent; involucre 10-12 mm. in diameter; bracts 2-seriate, linear-lanceolate, acute, purple-margined, sparse- and somewhat appressed-pubescent, 4-5 mm. long; ray- flowers about 30, 1 cm. long; achenes sparsely pubescent with glochidiate or straight and slightly emarginate hairs; pappus an inconspicuous whitish ring-like crown. Distribution: western Chihuahua, southward along mountains in the state of Durango, Mexico. [Vol. 20 32 ANNALS OF THE MISSOURI BOTANICAL GARDEN Specimens examined: Chihuahua: Guachachic, June 25, 1892, Hartman 523 (G, type); vicinity of Madera, alt. 2250 m., May 27-June 3, 1908, Edward Palmer 287 (M). Duuanuo: City of Durango and vicinity, Apr.-Nov. 1896, Edward Palmer 163 (M); Otinapa, July 25- Aug. 5, 1906, Edward Palmer 425 (M). 4. Astranthium mexicanum (Gray) Larsen, n. comb. Bdlis mexicana Gray, Smiths. Contr. [PI. Wright, pt. 1] 3: 93. 1852; Gray in Hemsl. Biol. Cent.-Am. Bot. 2: 118. 1881. Perennial from a branched ligneous base; stems 15-00 cm. high, several, branched above, striate, pubescent with spreading hairs; leaves appressed-pubescent, those of the stem sessile, entire, or saliently dentate, oblong-linear, gradually reduced toward the inflorescence, basal leaves ellipsoid-spathulate, crenate-dentate, 5-20 cm. long, .8-2.5 cm. broad; involucre .8- 1.4 cm. in diameter; bracts 2-3-seriate, cinereous-pubescent, linear-lanceolate, purple-tipped; ray-flowers white, numerous, .5-1.0 cm. long; pappus an inconspicuous ring-like crown or almost obsolete; achenes compressed, pubescent. Distribution: mountains of south-central Mexico. Specimens examined: Hidai.wo: Sierra de Pachuca, alt. 3076 m., Aug. 26, 1902, Pringle 9857 (M). Mexico: Apr. 26, 1849, Uregg 701 (M, type); along brooks, Ixtaccihuatl, March- July I 903, 1'urpus 169 (M); moist open woods and meadows about timber line, Ixtaccihuatl, Oct. 1905, Purpus 1576 (M); "above timber line," Popocatepetl, Oct. 1908, I'urpm 3640 (M). Federal District: Cima, Aug. 24, 1910, Orcutt 3768 (M); Serrania de Ajusco, alt. 3076 m., Aug. 8, 1896, Pringle 6442 (M). Mo hum: Toro, alt. 3015 m., Aug. 5, 1924, Fisher (M, No. 914805). Oaxaca: Sierra de San Felipe, alt. 3076 m., June 28, 1894, Pringle 4719 (M); Sierra de San Felipe, alt. 3076 m., Sept. 1894, Smith 261 (M). 4a. Var. chihuahuense Larsen, n. var. 22 Plate 3. Resembles the species in habit but differs from it in that it is much less pubescent; the leaf margins are essentially entire, ciliate; the pappus is more conspicuous than that of the species and consists of a narrow fluted crown; the achenes are glabrous, or glabrate. 22 Astranthium mexicanum (Gray) Larsen var. chihuahuense Larsen, n. var., plant* herbftcea perennis, 6-7 dm. alta, supra ramosa; marginibus foliorum integris vel subintegris; pappo ooroniforme, margine irregulariter sinuato-dentato; achaeniis glabris v«-l glabratis— Mkxico: canyons of Sierra Mad re, State of Chihuahua, Oct. 4, 1888, Pringle 2015 (M, No. 122921, type). 1933] LARSEN — ASTRANTHIUM AND RELATED GENERA 33 Distribution: State of Chihuahua, Mexico, known only from the type. Specimens examined: Chihuahua: canyons of Sierra Madre, Oct. 4, 1888, Pringle 2015 (M, No. 122921, type). 5, Astranthium xanthocomoides (Less.) Larsen, n. comb. Brachycome xanthocomoides Less. Syn. 192. 1832, nomen sub- nudum; Schlecht. in Linnaea 9: 265. 1835. Brachycome xeranthemoides Steud. Nomencl. Bot. ed. 2, 220. 1840, in part (typographical error). Keerlia linearifolia DC. Prodr. 5: 310. 1836, in part as to synon. Bellis xanthocomoides (Less.) Gray in Hemsl. Biol. Cent.-Am. Bot. 2: 118. 1881. Annual ; stems several, striate, procumbent or ascending, branching, leafy throughout, producing slender runners; leaves spathulate, apiculate, sparsely pubescent, gradually reduced toward the peduncles, 1.5-2.5 cm. long, .3-.8 cm. broad; pe- duncles 1-6 cm. long, densely pubescent at the base of the invo- lucre; involucre .8-1.2 cm. in diameter; bracts 2-seriate, lanceo- late, acute, pubescent along the lower portion of the main axis, membranaceous-margined ; ray-flowers white, .5-8 cm. long, about 16; pappus practically lacking; achenes compressed, pubescent, becoming more or less glabrate. Distribution: alpine meadows and open woods of east-central Mexico. Specimens examined: Vera Cruz: prope la Jaya (La Hoya), June 29, Schiede 206 (M, type). Hidalgo: meadows of Sierra de Pachuca, alt. 2760 m., July 17-28, 1898, Pringle 6888 (M); alpine meadows, Sierra de Pachuca, alt. 2923 m., Aug. 26, 1902, Pringle 9858 (M); moist meadows and open woods, Pachuca, July 1905, Purpus 1344 (M). 6. Astranthium purpurascens (Rob.) Larsen, n. comb. Bellis purpurascens Rob. Proc. Am. Acad. 27: 172. 1892. Perennial; roots of numerous fibers; stems erect, branched, 4- 45 cm. high, striate, pubescent with spreading hairs; leaves apiculate, pubescent with spreading hairs, margins somewhat ciliate; lower leaves oblong-ovate, 2.5-4.0 cm. long, 1.5-6 cm. broad ; upper leaves linear to linear-lanceolate, gradually reduced, 1 cm. or less in length at the base of the peduncles; peduncles 3-8 cm. long; involucre .8-1.0 cm. in diameter; involucral bracts about 14-16, lanceolate, acute, scarious-margined, only slightly 34 ANNALS OF THE MISSOURI BOTANICAL GARDEN I Vol. 20 ciliate near the apex, pubescent along the main axis with a few upwardly appressed hairs; rays 10-15, whitish-purple, .5-8 cm. long; pappus lacking; achenes glabrous, golden-brown at matu- rity. This species is closely allied to A. integrifolium (Michx.) Nutt.; the chief difference is to be found in achenial characters. Astran- thium integrifolium has achenes which are pubescent with glo- chidiate-tipped hairs whereas A. purpurascens (Rob.) Larsen has glabrous achenes. Distribution: known only from the type locality and Chiapas, Mexico. Specimens examined: San Luis Potosi: shaded grassy slopes, barranca of Las Canoas, Aug. 18, 1891, Pringle 3819 (M, cotype). Chiapas: without definite locality, coll. of 1864-70, Ghiesbreght 548 (M). 7. Astranthium integrifolium (Michx.) Nutt. Trans. Am. Phil. Soc. N. S. 7:312. 1841. Bellis integrifolia Michx. Fl. Bor. Am. 2: 131. 1803; Hook. Bot. Mag. 52 : pi. 3^55 of Brachycome xanthocomoides Less.; DC. Prodr. 5: 304. 1836; Raf. New Fl Am. 2: 24. 1836; Torr. & Gray, Fl. N. Am. 2: 189. 1842 Gray, Syn. Fl. N. Am. I 2 : 163. 1884, and ed. 2, 1886 and 1888 Britton & Brown, 111. Fl. 3 : 350. 1898, and ed. 2, 3 : 402. 1913 Britton, Manual, 943. 1901, and ed. 2, 1905; Rob. & Fern, in Gray, Manual, ed. 7, 799. 1908; Small, Fl. Southeastern U. S. 1202. 1903, and ed. 2, 1913. Bellis nutans Raf. New Fl. Am. 2: 23. 1836. Bellis parviflora Haf. Ibid. Eclipta integrifolia Spreng. Syst. Veg. 3: 602. 1826. Annual, openly branched from near the base, 8-45 cm. high, sparsely pubescent with spreading or subappressed hairs, fre- quently conspicuously pubescent near the base, leafy throughout; lower leaves oblong-spathulate, 2.5-4.0 cm. long, .8-1.4 cm. broad; stem-leaves oblong-ovate, sparsely pubescent, margins ciliate, 3.5 cm. long, 1.5 cm. broad, gradually reduced toward the base of the peduncle; peduncles 4-9 cm. long, densely pubescent at the base of the involucre; involucre 2-seriate, about .6-1.2 cm. in diameter, bracts lanceolate, acuminate, acute, membranous margins only slightly lacerate, sparsely pubescent along the main 1933] LARSEN — ASTRANTHIUM AND RELATED GENERA 35 axis with a few spreading hairs; ray-flowers fertile, 16-22, pur- plish-blue; achenes compressed, pubescent with glochidiate- tipped hairs. Distribution: sandy soil, south-central Kentucky to northwestern Georgia, west- ward to southeastern Kansas and eastern Oklahoma. Specimens examined: Georgia: dry ground, Ringold Road, Chickamauga, May 27, 1911, Churchill (M, No. 839094). Mississippi: Tchula, Holmes Co., Apr. 18, 1927, Woodson & Anderson 1555 (M). Kentucky: Bowling Green, May 6, 1892, Price (M, No. 122882). Tennessee: West Nashville, May 26-27, 1909, Eggleston 1+1+22 (M); moist field, Joelton, Davidson Co., July 16, 1922, Svenson 118 (G); copses around Nashville, May- June, Gattinger 1297 (M); rocky open hillsides, near Erin, Houston Co., May 24, 1920, E. J. Palmer 1762S (M, G). Missouri: gravelly barrens, Noel, May 10, 1915, Bush 7534 (M, G); gravelly barrens, Noel, May 10, 1915, Bush 7534 A (M); rocky open ground, near Jane, MacDonald Co., May 23, 1931, E. J. Palmer 39297 (M); rocky slopes, bald knobs, along creek, near Oasis, Taney Co., June 3, 1931, E. J. Palmer 39483 (M). Arkansas: river bottoms near Fayetteville, June 1835, Engelmann 129 [607] (M); alluvial soil, waste places, cultivated fields, Fayetteville, May, Harvey 45 (M); Fayetteville, 1880, Harvey (M, No. 122880); Hot Springs, May, 1879, Soulard (M, No. 122885); sandy soil, Cotter, Marion Co., June 15, 1914, E. J. Palmer 6992 (M). Kansas: rocky soil, Chautauqua Co., May 7, 1897, Hitchcock 1055 (M). Oklahoma: in woods and meadows, common near Tishomingo, Johnston Co., Apr. 27, 1916, Houghton 3549 (G); clay washes, near Ardmore, Carter Co., Apr. 17, 1913, Stevens 77 (M); Arbuckle Mts., Crusher, May 12, 1916, Emig 606 (M); Catoosa, May 8, 1895, Bush 897 (M); Ind. Terr., 1877, Butler (M, No. 122886); upland prairies, sandy soil, near Howe, Le Flore Co., May 25, 1931, E. J. Palmer 39343 (M). 7a. Var. ciliatum (Raf.) Larsen, n. comb. PL 4, fig. Bellis ciliata Raf. New. Fl. Am. 2: 24. 1836. Bellis integrifolia Gray, Smiths. Contr. [PI. Wright, pt. 5:78. 1853, in part. Stems several, diffusely branched from near the base, 1-3 dm. high, terminated by long slender peduncles; leaves oblong- spathulate to linear-oblong, reduced to small bract-like leaves at the base of the peduncles, .8-3.0 cm. long, .2-.8 cm. broad, sparsely pubescent with subappressed hairs, margins ciliate; peduncles 4-6 cm. long; involucre .5-8 cm. in diameter; in- volucral bracts lance-elliptic, acute, the narrow membranaceous margins lacerately ciliate; ray-flowers about 9-15; achenes pubes- cent with glochidiate-tipped hairs. Distribution: from southeastern Oklahoma and southwestern Arkansas to south- central Texas, west to the Pecos. [Vol. 20 3C> ANNALS OF THE MISSOURI BOTANICAL GARDEN Specimens examined: Arkansas: moist open ground, Fayetteville, Washington Co., July 6, 1915, E. J. Palmer 8176 (M); Coal Bank, May 14, 1895, Bush 895 (M); Redfork, May 14, 1895, Bmh 896 (M). Oklahoma: open woods near Idabel, McCurtain Co., May 18, 1916, Houghton 8645 (C). Tex ah: Apr. May, 1844, Limiheimer 25 1 (M); "Pine's Island," May 5, 1903, Rever- sion (M, No. 122889); Rrverchon 440 in part (M, No. 122890); clay barrens, Bryan, Brazos Co., May 27, 1915, E. J. Palmer 7785 (M); Pecos and Limpio, June 1851-2, Wright 1176 in part (M, No. 123093); dry banks, Hempstead, Apr. 24, 1872, Hall 806 (M). 7b. Var. rosulatum Larsen, n. var. 23 PL 4, fig. 2. Belli* integrifolia Gray, Smiths. Contr. [PI. Wright, pt. 2] 5: 78. 1853, in part. Stems several, sparsely branched near the base, pubescent with spreading hairs, .5-1.5 dm. high; leaves sparsely pubescent with subvillose hairs; basal leaves numerous, forming a rosette, oblong- spathulate, 2-7 cm. long, .5-1 cm. broad; stem-leaves oblong- spathulate to linear-lanceolate, gradually reduced to the base of the peduncle; peduncle 1.5-5 cm. long; involucre about .7 cm. in diameter; bracts lance-elliptic, acute, the narrow membranaceous margins somewhat lacerately ciliate near the apex; ray-flowers about 16; achenes densely pubescent with glochidiate-tipped hairs. Distribution: central Oklahoma and eastern Texas west to Presidio County. Specimens examined: ( )ki,aiioma: open woods near Mannsville, Johnston Co., May 16, 1916, Griffith 8455 (M, G); Limestone Cap, Apr. 16, 1877, Butler 63 (M); Sapulpa, Apr. 28, 1895, Bush 927 (M); "Arkansas," May 20, 1895, Bruh 929 (M). Texas: sandy prairies, Columbia, Brazoria Co., March 25, 1914, E. J. Palmer 6028 (M); Victoria, Victoria Co., March 5, 1916, E. J. Palmer 9070 (M); sandy open ground, Larissa, Cherokee Co., Apr. 7, 1916, E. J. Palmer 9388 (M); sandy prairies, Matagorda, Matagorda Co., March 5, 19U,E.J. Palmer 4855 (M, No. 753171, type); on plains, Handley, Apr. 15, 1913, Ruth 74 (M, G); along railroads near Houston, Apr. 23, 1899, Eggert (M, Nob. 122893 & 122895); Terrell, Apr. 6, 1903, Reverchon 4006 (M); prairies near Victoria, Apr. 7 & 10, 1900, Eggert (M, No. 122899); Pecos and Limpio, June 1851-2, Wright 1176 in part (M, No. 123091); near San Felipe, " Astranthium integrifolium (Michx.) Nutt. var. rosulatum Larsen, n. var., caule simplice vel basi ramoso, 0.5-1.5 dm. alto, subvilloso; foliis basalibus rosulatis, oblongo-npathulatis, 2-7 cm. longis, usque ad 1 cm. latis, integris, ad apicem rotun- datis vel obtusis, sparse pubescentibus; foliis caulinis oblongo-spathulatis vel lineari- lanceolatis, integris, Hiiperne sensim minoribus; pedunculis 1-6 cm. longis. — Texas: on sandy prairies, Matagorda, Matagorda County, March 5, 1914, E. J. Palmer 4855 (M, 753171, type). 1933] LARSEN — ASTRANTHIUM AND RELATED GENERA 37 Apr. 1839, Lindheimer (M, No. 122904); along river, Columbia, Apr. 19, 1899, Bush 195 (M); on prairie, Chenango Junction, Apr. 18, 1900, Bush 60 (M); Reverchon 440 in part (M, No. 122898); open ground, along railway grade near Marfa, Presidio Co., June 18, 1926, E. J. Palmer 31086 (M). KEERLIA Keerlia Gray, Bost. Jour. Nat. Hist. [PI. Lindh. pt. 2] 6: 221, 222. 1850, in part as to K. bellidifolia and K. effusa, not DC; Smiths. Contr. [PL Wright, pt. 1] 3: 92. 1852, not DC; Syn. Fl. N. Am. I 2 : 164. 1884, and ed. 2, 164. 1886 and 1888. Bourdonia Greene, Erythea 1 : 207. 1893. Herbaceous, caulescent, pubescent annuals or perennials. Leaves alternate, sessile or petioled, oblong to obovate-spathulate, entire. Involucre of imbricated lanceolate bracts, with mem- branaceous margins. Heads few-flowered. Ray-flowers in a single series, pistillate, fertile. Disk-flowers tubular, five-lobed, frequently sterile. Branches of the style lanceolate, acutish, hairy toward the tip. Pappus a thickened ring-like or slightly lacerated crown. Achenes compressed, glabrate or hirsute- pubescent. Type species: K. bellidifolia Gray & Engelm. Proc. Am. Acad. 1 : 47. 1848, which was based on No. 415 of Lindheimer, collected above Guadeloupe, Texas, 1845-1846. Key to the Species Stems branched from the base 8. K. bellidifolia. Stems simple to the inflorescence 9. K. effusa 8. Keerlia bellidifolia Gray and Engelm. Proc. Am. Acad. 1 : 47. 1848; Gray, Bost. Jour. Nat. Hist. [PL Lindh. pt. 2] 6: 220. 1850; Smiths. Contr. [PI. Wright, pt. 1] 3: 92. 1852; Syn. Fl. N. Am. I 2 : 164. 1884, and ed. 2, 164. 1886 and 1888. Bourdonia bellidifolia (Gray & Engelm.) Greene, Erythea 1 : 207. 1893. Annual; the caudex giving rise to much-branched stems 9-30 cm. high, leafy throughout, striate, pubescent with multicellular spreading hairs; leaves thin, sparsely pubescent, apiculate, lower leaves obovate-spathulate, uppermost somewhat linear, gradually reduced toward the inflorescence, 1-4 cm. long, .4-1.0 cm. broad, narrowed below to a subpetiolate base; heads small; involucre [Vol.. 20 38 ANNALS OF THE MISSOURI BOTANICAL GARDEN .3-.5 cm. in diameter, bracts 2-3-seriate, glabrous, linear-lanceo- late, acute, with membranaceous margins; ray-flowers 4-15, blue, fertile; disk-flowers 15-20, frequently sterile, style branches shorter than those of the ray-flowers; pappus a thickened ring- like crown; achenes nerved, compressed, glabrate. Distnl.ution: southern Texas; doubtless also in adjacent Mexico. Specimens examined: Texah: along Nueces Bay, Nueces Co., alt. 6 in., March 12, 1894, Heller 1436 (M); Rock Springs, Apr. 17, 1930, Marcus E. Jones 26575 (M); above Guada- lupe, 1845-46, Lindheimer 415 (M, co-type); near New Braunfels, May 1848, Lindheimer 628 (M); Comanche Spring, New Braunfels, Apr. 1850, Lindheimer 982 (M); rocky calcareous ground, Del Rio, Valverde Co., March 26, 1917, E. J. Palmer 11384 (M); dry limestone hills, Concan, Uvalde Co., Apr. 13, 1917, E. J. Palmer 11550 (M); vicinity of San Antonio, Apr. 1919, von Schrenk (M, No. 88063); near Austin, March 17, 1908, York 410 (M). 9. Keerlia effusa Gray, Bost. Jour. Nat. Hist. [PI. Lindh. pt. 2] 6: 222. 1850; Smiths. Contr. [PI. Wright, pt. 1] 3 : 93. 1852; Syn. Fl. N. Am. I 2 : 1G5. 1884, and ed. 2, 165. 1886 and 1888. Bourdonia effusa (Gray) Greene, Erythea 1 : 207. 1893. Perennial, 15-60 cm. high ; stem simple below, diffusely branched above into an open glabrous panicle; stem and leaves hispid- pubescent; leaves thin, lowermost leaves obovate-spathulate, 3-9 cm. long, .5-1.3 cm. broad, narrowed at the base into a petiole, those of the stem oblong with a broad, sessile, somewhat amplexicaul base, gradually reduced toward the inflorescence; heads numerous, small; involucre turbinate, 2-3 cm. in diameter; involucral bracts glabrous, 3-4 acute, membranaceous-margined; ray-flowers 4-7, white, fertile; disk flowers 6-9, sterile; pappus consisting of a thickened irregularis lacerate crown; achenes compressed, hispid-pubescent. Distribution: south-central Texas. Specimens examined: 499 1847-48 Lindheimer 629 (M, co-type); dry hillside thickets, Leakey, Edwards Co., June 10, 1916, E. ./. Palmer 10166 (M); dry hillside thickets, Spanish l'ass, Kendall Co., Sept. 28, 1916, E. J. Palmer 10839 (M); San Geronimo Creek, June 1884, Reverchon 1534 (M); Cunanche Spring, New IJraunfels, 18-19, Lindheimer 9SS (M). 1933] LARSEN — ASTRANTHITJM AND RELATED GENERA 39 List of Exsiccatae The distribution numbers are printed in italics. The number in parenthesis is the species number used in this revision. Berlandier, J. L. 499 (9). F Bush, B. F. 897, 7534, 75S4A (7); 895, 896 (7a); 60, 195, 927, 929 (7b). Butler, G. D— (7); 68 (7b). Churchill, J. R. (7). Eggert, H. (7b). Eggleston, W. W. 4422 (7). Emig, W. H. 606 (7). Engelmann, G. 129 [607] (7). Fisher, G. L. (4). Gattinger, A. 1297 (7). Ghiesbreght, A. 548 (6). Gregg, J. 701 (4) . Griffith, F. 3455 (7b). Hall, Elihu. 806 (7a). Hartman, C. V. 523 (3). Harvey, F. L. — , 45 (7). Heller, A. A. 1436 (8). Hitchcock, A. S. 1055 (7). Houghton, H. W. 3549 (7); 3645 (7a). Jermy, Gustav. 815 (9). Jones, Marcus E. 26575 (8). Lindheimer, F. 251 (7a); — , (7b); 416, 628, 982 (8); 629, 938 (9). Nelson, E. W. 4786 (1). Orcutt, C. R. 3768 (4). Palmer, E. J. 5992, 1 7623, 89297, 39348, 89483(7); 7785, 81 76 (7a) ; 4856, 5028, 9070, 9388, 31036 (7b); 11384, 11560 (8); 10166, 10839 (9). Palmer, Edward, 146 (2); 163, 287, 425 (3). Price, Sadie F. (7). Pringle, C. G. 4412 (2); 4719, 6442, 9857 (4); 2015 (4a); 6888, 9858 (5); 8819 (6). ', 1575, 3640 (4) ; 1344 (5). Reverchon, J. — , 440 in part (7a); (7b); 4006, 440 in part (7b); 1534 Ruth, Albert. 74 (7b). Schiede, G. 206 (5). Smith, Charles L. 261 (4). Soulard, Mary. — (7). Stevens, G. W. 77 (7). Svenson, H. K. 118 (7). von Schrenk, H. (8). Woodson, R. E. Jr., and Anderson, E 1555 (7). Wright, Charles. 1176 in part (7a); 1. in part (7b). York, H. H. 410 (8). [Vol. 20. 1933] 40 ANNALS OF THE MISSOURI BOTANICAL GARDEN PLATE 2 Astranthium integrifolium (Michx.) Nutt. from Egglesion 4422 in the Missouri Botanical Garden Herbarium. Pig Fig Fig Fig Fig Fig 1. Involucre and receptacle, X 5. 2. Ray-flower, X 10. 3. Disk-flower, X 10. 4. Stamen, X 10. 5. Style-branches of the disk -flower, X 10. 6. Style-branch greatly enlarged. Belli s perennis L. from Pring, in the Missouri Botanical Garden Herbarium No. 918135. Fig. 7. Involucre and receptacle, X 5. Fig. 8. Ray-flow X 10. Fig. 9. Disk-flower, X 10. Fig. 10. Stamen, X 10. Fig. 11. Style-branches of the disk-flower, X 10. Fig. 12. Style-branch greatly enlarged. Ann. Mo- Bot. Card., Vol. 20, 1933 Plate 2 // u / ( v IT 5 10 LARSEN— ASTRA XTHIUM VXD RELATED GENERA [Vol. 20, 1933] 42 ANNALS OF THE MISSOURI BOTANICAL GARDEN PLATE 3 Idtranthium mexicanum (Gray) Larsen var. chihuahncnse Larsen. From the type specimen, Pringle 2015, in the Missouri Botanical Garden Herbarium. Ann. Mo. Hot. Gaki>., Vol. 20, 1933 Plate 3 C« G PHZ.KOf \ Nftft. itah :? OBrntussA 2015 Aphantostephus Arkansanus, Gray. % ■ *>!. V. . ji J C^* ; * ■ > -* # '" » ** LARSEN— ASTRANTIIIL'M AND RELATED GEXERA [Vol. 20, 1933] 44 ANNALS OF THE MISSOURI BOTANICAL GARDEN PLATE 4 Fig. 1. Aslranthium integrij olium (Michx.) Nutt. var. ciliatum (Raf.) Larsen. From Lindheimer £51 y in the Missouri Botanical Garden Herbarium. Fig. 2. Astranthiuvi intcgrifolium (Michx.) Nutt. var. rosulatum Larsen. From the type specimen, Palmer 4855, in the Missouri Botanical Garden Herbarium. Ann. Mo. Hot. Gakd., Vol. 20, l ( .>:i3 Plate 4 tr 1 > > H I 1 —4 It * V B SB I I - t 1 X s % * ■'. 3 . 7 > • c to i I • • . .- ■ . FERNS AND FERN ALLIES OF MISSOURI 1 M. ELIZABETH PINKERTON Assistant in Botany, Henry Shaw School of Botany of Washington University This paper is based primarily upon material in the herbarium of the Missouri Botanical Garden, the University of Missouri, and the private collections of Mr. John H. Kellogg and of the author. After each specific name reference is given to the original publication; this is followed by reference to the two standard manuals and to an illustrated monograph covering the fern-flora of the region concerned, where the species or variety is treated under the same or a different name. These are: Gray, 'New Manual of Botany/ ed. 7, 1908; Britton and Brown, 'Illustrated Flora of the Northern States and Canada/ ed. 1, 1896, and ed. 2, 1913; and Eaton, 'Ferns of North America/ vol. 1, 1879 (plates 1-45) and vol. 2, 1880 (plates 46-81). Time of fruiting, the general distribution for North America, and the habitat for Missouri in particular precede the specific description. The names of the principal collectors whose material was ex- amined are listed, and definite citations made when the plant is rare. Key to Families 1. Plants with short vertical stems or rootstocks 2 1. Plants with horizontally spreading stems or rootstocks 4 2. Leaves onion-like, producing sporangia at their bases Isoetaceae 2. Leaves neither onion-like nor producing sporangia at their bases 3 3. Sporangia borne on under sides of leaves Polypodiaceae 3. Sporangia borne terminally in special fruiting clusters Ophioglossaceae 4. Sterile leaves large, usually compound or variously divided 5 4. Sterile leaves small, often scale-like 6 5. Fertile segments leaf -like or pod-like; if the latter, sporangia covered with indusia Polypodiaceae 5. Fertile segments not leaf -like; sporangia naked Osmundaceae 6. Plants truly aquatic Salviniaceae 6. Plants terrestrial 7 7. Stems jointed; sporangia clustered underneath the scales of terminal cone- like spikes Equisetaceae 7. Stems not jointed; sporangia borne on upper surfaces of leaves 8 1 This article is a portion only of a thesis submitted to the Board of Graduate Studies of Washington University in partial fulfillment for the requirements of the degree of master of science. Issued April 29, 1933. Ann. Mo. Bot. Gard., Vol. 20, 1933. (45) 46 [Vol. 20 ANNALS OF THE MISSOURI BOTANICAL GARDEN 8. Spores all of the same kind Lycopodi 8. Spores of two kinds (larger macrospores and much smaller microspores).. . taceae Selaginellaceae Ophioglossaceae 1. Veins free; sporangia separate in compound inflorescences Botrychium 1. Veins reticulate; sporangia disposed in a solitary spike Ophioglossum Fig. 1. Key map to distribution chart (tig. 2) showing location of counties in Missouri. OPHIOGLOSSUM 1. Sterile segments obtuse, usually solitary; middle areoles long and narrow, outer hexagonal and containing a few included veinlets 1. 0. vulgatum 1. Sterile segments cuspidate, 2-5; areoles wide and containing many anasto- mosing veinlets 2.0. Engelmanni 1. Ophioglossum vulgatum L. Sp. PL 2 : 1062. 1753. Gray, p 47; Brit I on & Brown, ed. 1 and 2, 1:2. 1 2 3 ' 4 5 6 7 8 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 it ?! Adiantum CaDillus-Veneria " oedatum AaDlanlum Bradley! cryptolepia ebenoidea " plnnatifidum " platyneuron " resiliens " Trlohomanea Athyrium acroatichoidea n anguatlfolium " anguatum H aaplanloidaa Azolla carollnlana ootrychium obliquum " " diss actum " " tenuifoliua " virglnlanura Camptoaorua rhizophyllua (jhellanthea alabamenala " Feel " lanoaa Cyatopterla bulblfora H fraffllla Dennataedtia ounctilobula Eauiaetum arvense " laevi^atum praealtum Butleri Engelmanni melanopoda Lycopodium lucidulum " complanatum flabelllformej Notholaena dealbata Onoclea aenaibilla UDhiogloaaum rjnp elmanni " vulKatum Oamunda clnnaroomea " Claytonlana " regalia BDectabilla Pellaaa atropurDurea tt glabella Polypodlum polyoodioidea " vlrginianum Polyatichum acrostlcholdes Pteretls noduloaa Pterldium latiuaculum oelactlnella aooda " rupeatrla Thelypteris haxagonoptera ** marginalia " paluatria pubescena apinuloaa Intermedia Woodala obtusa i.oodwardia areolata CO 25 bd COO OS MM 3§§ o .J a 3o « o a- ^J 33 O E-« CO co« MO §2 H w CO»"» i Q Was cm 2se to w 25 c _ • as >ju £ t-« w o< o CO Ou. OQ O CO s o o CO a. Q O.M MQC co q co 3 o M S CO CO CO _ w a. + 4-4-44 4-4 4- 4 4-4 + + 4- 4 4 + 444 44 44 4 + 4 4 4 4 4 4 4 4444 4444 + + t 4 444 [444444 4- 4 isoetes N N 4- + -f-4-t 444 444 44 4444 4 44 4 44 4 4 4- 4 4 4-4- » 4 4 44 4-4 + 4 44444 44 4-4 4- 4 4"44 + 44 4-4-4 444 44-44 4 4 4-44 44 +4-4 it 4" 4 4 4-4-4- + + 4 44 + t 4 4- 4 + 44 4 4 4 4 4- 4 + 4 4 & K J O O Id <3 w m =* 025 o < < c a: os j e- D < O < < o at u M < H 25 CO CO Wo yy cu o z * o o o ►J *t M oBph rn ** co »-3 o 3 M Q ccw M o- Sh C5 3 0,M 3 * 025 3- O OO o k oa« 25 25 < W o Q a: a a o 3^ W 25 J O o 2 3iS M w o ex w 25 o f » K J W O tc. K M 33S§ CO 25 05 CO M O 0< Oh) OS w « CO h4 cq o w 4 OaOM'OCOfeCQEQ A,^ w w OS o < t J H OO) J ev coco w 01 >i M W ^ h tx3 o: m 5 OS 2 X > »J ot^a q O CO o oos^,^ 25 3 O OS • Q H W CO O 25 >m CQ W >m W 3S OS W OS OS 25 E-< E- Q ^ X O OS 25 _ 3 ^ ^fc 2 w y ^h3^ < 25 25 O O 4 a X0S 25 s < CO o o w o o ^ IxJ 25 E-* CU E^ Q < 5 w 52 S o o « > cq»-> 25 a a < o * 35 W CO W < H M os as h — h flO765. 2. Ophioglossum Engelmanni Prantl in Ber. Deut. Bot. Ges. 1 : 351. 1883. Gray, p. 47; Britton & Brown, ed. 1 and 2, 1:2. Rootstock cylindrical; common stalk mostly below ground, sheathed by persistent leaf-bases; sterile segments sessile, ellip- tic to ovate, 1-3.5 inches long, 0.5-2 inches broad; basal veins 13 or more, transverse ones oblique; sporophyll 0.5-1 inch long, apiculate. March to October. Distribution : South Carolina to Florida, westward to Arizona. Limestone glades, especially where there is a thin layer of fine rich and damp soil. Specimens examined: Palmer, Kellogg, Letterman, Engelmann, Bush, Pinkerton, Eggert f Greenman. BOTRYCHIUM 1. Bud wholly enclosed; cells of epidermis straight; sterile blade petiolate and never over 7 inches broad; common stalk mostly underground 2 1. Bud exposed along one side at base of rootstock; cells of epidermis flexuous; sterile blade sessile, 2-16 inches broad and nearly as long; common stalk above ground, half the length of plant 4 2. Ultimate segments deeply laciniate 2. B. obliquum var. dissectum 2. Ultimate segments serrulate-dentate 3 3. Plant coarse; common stalk very short, underground; frond membranaceous on drying 1. B. obliquum 3. Plant slender; common stalk up to 2 inches above ground; frond coriaceous on drying 8. B. obliquum var. tenuifolium 4. Ultimate segments spatulate-shaped; ripe sporangia straw-colored, opening but slightly in dehiscence 5. B. virginianum var. intermedium 4. Ultimate segments acute; ripe sporangia brown, opening widely in dehiscence. 4. B. virginianum 1. Botrychium obliquum Miihl. in Willd. Sp. PL 5: 63. 1810. Gray, p. 48; Britton & Brown, ed. 1, 1: 3, ed. 2, 1: 5; Eaton, pi. 20, fig. 2. B. ternatum Sw. in Schrad. Jour, fur die Bot. 2 (1800): 111. 1801. B. ternatum var. obliquum D. C. Eaton, Ferns N. Am. 1: 149. 1879. 48 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 20 Plant 5-15 inches tall; bud pilose; sterile blade long-stalked, 2-5 inches broad, tripinnatifid or tripinnate below; ultimate segments obliquely ovate or oblong-lanceolate, acutish terminal ones elongate; sporophyll long-stalked, usually stout, several- pinnate. August to October. Distribution: Maine to Alabama, westward to Michigan and Texas. Moist woods, neutral soil. SpeciniciiH examined: Palmer, Bush, Kellogg, Letterman, Eggert, Trelease. 2. Botrychium obliquum var. dissectum Prantl in K. Bot. Gard. Berlin, Jahrb. 3: 342. 1884. Gray, p. 49; Britton & Brown, ed. 1, 1 : 3, ed. 2, 1:5; Eaton, pi. 20, fig. 1. B. dissectum Spreng. Anleit. 3: 172. 1804. B. ternatum var. dissectum D. C. Eaton, Ferns N. Am. 1: 150. 1879. Character of rootstock, fruiting stalks, and texture of the plant all similar to the species; frond subternately divided, basal divisions unequally and broadly deltoid, decompound; the upper and secondary pinnae deltoid-lanceolate, pinnate, with laciniate the ultimate divisi d Distribution: Maine to Florida, westward to Illinois and Missouri. Rich moist woods, deeply shaded. Specimens examined: Eggert, Letterman, Pinkerton. 3. Botrychium obliquum var. tenuifolium (Underw.) Gilbert in Fern Bull. 11 : 99. 1903. Gray, p. 49. PI. 5, fig. 2. Plant small, up to 12 inches high; common stalk up to 2 inches high, not all underground, slender; blade ternate with few divi- sions, 5-10 cm. broad and about 5 cm. long; leaves coriaceous; ultimate segments broad, about 1.5 cm. long, ovate, acutish, conspicuously serrulate. September and October. Distribution: Virginia to Arkansas. Southern Missouri — swampy land. Specimens examined: Dunklin Co., Bush 134, and coll. of Sept. 28 and Oct. 28, 1897, Trelease; Butler Co., Bush SI 10. 4. Botrychium virginianum (L.) Sw. in Schrad. Jour, ftir die Bot. 2 (1800) : 111. 1801. Gray, p. 49; Britton & Brown, ed. 1, 1 : 4, ed. 2, 1:6; Eaton, pi. 33. Common stalk slender, above ground 1/2-2/3 the length of 1933) PINKERTON — FERNS OF MISSOURI 49 plant; blade sessile or nearly so, membranaceous, ultimate segments toothed; sporophyll long-stalked, bi-tripinnate. June and July. Distribution: New Brunswick to Alabama, westward to Idaho and Texas. Rich moist woods. Specimens examined: Davis, Engelmann, Letterman, Palmer , Bush, Pinkerton, Eggert, Kellogg, Trelease, Larsen, Daniels, Emig, Blankenship. 5. Botrychium virginianum var. intermedium Butters in Rho- dora 19:207. 1917. Ultimate segments of the sterile frond spatulate, penultimate ones ovate, not crowded; segments of the fertile frond opening wide in dehiscence; sporangia straw-colored, up to 0.8 mm. long. Specimens examined: Monteer, Shannon Co., Bush 4721+; Whiteside, Lincoln Co., Davis, coll. of 1910. The writer feels that the species is so generally variable that this variety is not very distinct. POLYPODIACEAE 1. Indusia lacking 2 1. Indusia present, at least in early stages 4 2. Fruit dots near margin, becoming confluent, somewhat protected by over- lapping leaf margins; lower surfaces of leaves covered with a white powder. Notholaena 2. Fruit dots separate, round, absolutely unprotected 3 3. Fronds linear, once-pinnatifid Polypodium 3. Fronds triangular, more than once-pinnatifid Thelypteris 4. Fronds dimorphic (fertile fronds not leaf -like) 5 4. Fronds monomorphic 6 5. Sterile fronds pinnatifid; veins anastomosed; fertile segments bipinnate . Onoclea 5. Sterile fronds bipinnatifid; veins free and unbranched; fertile segments pinnate Pteretis 6. Indusia marginal 7 6. Indusia not marginal 11 7. Indusia not formed of revolute margins, but cup-shaped and opening ter- minally Dennstaedtia 7. Indusia formed in part at least by revolute margins 8 8. Indusia definitely interrupted, occurring in separate little rounded flaps. . . . Adiantum 8. Indusia continuous or only slightly broken 9 9. Indusia double Pteridium 9. Indusia single 10 10. Blades with a few large, relatively smooth segments Pellaea 10. Blades with many small, usually tomentose or hairy segments Cheilanthes 11. Sori elongated 12 11. Sori rounded 15 12. Sori dispersed in two rows parallel to the midrib Woodwardia 50 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 20 12. Sori dispersed irregularly on lateral veins 13 13. Veins anastomosed Camptosorus 13. Veins free 14 14. Sori straight Asplenium 14. Sori mostly curved over the end of veins Athyrium 15. Indusia peltate 16 15. Indusia attached at base or at one side 17 16. Indusia orbicular, attached centrally Polystichum 16. Induiia reniform, attached at sinus Thelypteris 17. Plan I s never glandular, but scaly on stipes and rootstocks; segments more or less acute; indusia tapering, attached at one side and becoming obscure at maturity Cystopleris 17. Plants glandular, also scaly; segments obtuse; indusia entire when young, splitting at top into several stellate segments Woodsia ADIANTUM 1. Main rachis unbranched LA. Capillus-Veneris 1. Main rachis dichotomously branched with 4-5 pinnae on each side. .£. A. pedatum 1. Adiantum Capillus-Veneris L. Sp. PL 2: 1096. 1753. Gray, p. 36; Britton & Brown, ed. 1, 1: 27, ed. 2, 2: 31; Eaton, pi 87. Venus hair. Rootstock creeping, chaffy; stipe very slender, 3-12 inches long, black or brownish, slightly scaly; frond ovate-lanceolate, 6-12 inches long, 4-12 inches wide at base; pinnules wedge- obovate or rhomboid, long-stalked, glabrous, membranaceous, margins variously incised, veinlets flabellately forking from base; fruit dots lunate or transversely oblong. June to August. Distribution: New Jersey to Florida, westward to South Dakota and California. Moist rocky places, ravines, wet limestone cliffs. Specimens examined: Kellogg, Daniels, Bush, Palmer, Trelease, Emig, Shepard. 2. Adiantum pedatum L. Sp. PL 2: 1095. 1753. Gray, p. 35; Britton & Brown, ed. 1, 1: 27, ed. 2, 2: 31; Eaton, pi. 18. Maidenhair. Rootstock long, creeping, chaffy; stipe 9-18 inches long, shining, dark brown to black, slightly scaly at base, once-forked at summit, each division bearing on one side only several pinnate divisions (occasionally tri-forked); blade reniform-orbicular, 8-18 inches broad, membranaceous, glabrous; pinnules short- stalked, oblong, triangular or end ones fan-shaped; lower margin entire, all veins proceeding from it, upper margin lobed. June to August. 1933] PINKERTON — FERNS OF MISSOURI 51 Distribution: Nova Scotia and Quebec to Georgia, westward to Alaska and California. Moist rich woods. Specimens examined: Bush, Davis, Engelmann, Palmer, Trelease, Pinkerton, Woodson, Kellogg, Broadhead, Emig, Mackenzie. ASPLENIUM 1. Blades pinnatifid, or only lower segments pinnate; apices long-attenuate 2 1. Blades 1-3-pinnate; apices not long-attenuate 3 2. Blades membranaceous; lower midribs black and shining on under side; apices crenate, sometimes proliferous 3. A. ebenoides 2. Blades subcoriaceous; midribs green, herbaceous; prolongations sinuous- margined, not proliferous 1+. A. pinnatifidum 3. Blades 1-pinnate only; pinnae of regular shape; margins not deeply dissected. 4 3. Blades 2-3-pinnate or -pinnatifid; pinnae of irregular shape; margins deeply cut 6 4. Pinnae mostly roundish, not auriculate 7. A. Trichomanes 4. Pinnae oblong or lanceolate, auriculate 5 5. Stipes and rachises shiny black, slender; pinnae mostly opposite; sori few and nearer margins than costae 6. A. resiliens 5. Stipes and rachises shiny reddish brown, coarse; pinnae mostly alternate; sori numerous, near costae 6. A. platyneuron 6. Stipes and rachises green throughout; ultimate segments few, cuneate; margins fimbriate 2. A. cryptolepis 6. Stipes and rachises shining chestnut-brown; lower pinnae divided into obtuse segments; margins crenate LA. Bradleyi 1. Asplenium Bradleyi D. C. Eaton, Bull. Torr. Bot. Club 4: 11. 1873. Gray, p. 39; Britton & Brown, ed. 1, 1 : 26, ed. 2, 1 : 30; Eaton, pi. 51, figs, 4-8. Rootstock short, covered with narrow acuminate blackish- fuscous scales; fronds 4-7 inches high, oblong-lanceolate; stipes 2-3.5 inches long, dark chestnut and shining, tufted, slender; rachises brown, or green above; pinnae numerous, lower ones no larger than the middle ones, obtuse or acutish, toothed, in the largest fronds pinnatifid into oblong lobes which are toothed at apices; sori short, borne near midveins, becoming confluent; indusia membranaceous, persistent. July to September. Distribution: New York to Georgia, westward to Missouri and Arkansas. On sandstone or chert outcrops — comparatively rare and local. Specimens examined: Palmer, Bash, Trelease, Mackenzie, Greene, Pinkerton, Van Dugen, Shepard. 2. Asplenium cryptolepis (L.) Fernald in Rhodora 30: 37. 52 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 20 1928. Gray, p. 39; Britton & Brown, ed. 1, 1: 25, ed. 2, 1: 29; Eaton, pi. 15, fig. 1. Asplenium Ruta-muraria L. Sp. PL 2: 1081. 1753. Rootstock short, creeping, entangled, tufted; fronds 1-6 inches tall; stipes and rachises entirely green or slightly brown at base; blades deltoid-ovate, smooth, subcoriaceous, bi-tri- pinnate; ultimate segments few, stalked, 3-14 mm. long, narrowly cuneate to roundish obovate; margins deeply fimbriate; veins flabellate, no midveins; sori few (2-4 per pinna), oblong, covering whole segment when mature; indusia delicate with ciliated margin. July to September. Distribution: Vermont to Georgia, westward to Illinois and Missouri. Shaded limestone cliffs — scarce Specimens examined: Treleasv, Palmer, Bush, Russell. 3. Asplenium ebenoides R. R. Scott, Berkeley in Roy. Hort. Soc. Jour. N. S. 1: 137. 1866. Gray, p. 38; Britton & Brown, ed. 1, 1: 23, ed. 2, 1: 26; Eaton, pi 4, fig. 2. Rootstock short, creeping, chaffy; fronds up to a foot high; stipes tufted, 4-9 inches long, young ones reddish brown, older ones black, shining, slender; lower rachises dark and shining underneath; blades firm-membranaceous, triangular-lanceolate, variable, 3-12 inches long, 1-3 inches at base, tapering to a long- acuminate apex which may become proliferous, lowest divisions distinct, shorter; sori numerous throughout, mostly single and opening obliquely upwards. August and September. Distribution: Vermont to Alabama, westward to Missouri. Limestone — rare. Specimens examined : Trelease, Russell. 4. Asplenium pinnatifidum Nutt. Gen. 2: 251. 1818. Gray, p. 38; Britton & Brown, ed. 1, 1: 22, ed. 2, 1: 27; Eaton, pi. 8, fig- 2. Rootstock short, creeping, branched, chaffy; fronds 6-9 inches high; stipes brownish near base and green above, clustered; blades 2-5 inches long, subcoriaceous, herbaceous, lanceolate- acuminate from broad and sub-hastate base, pinnatifid; the basal pinnae sometimes long-attenuate, lower lobes of pinnae roundish-ovate, margin crenate, upper pinnae gradually smaller and more adnate to winged rachises; sori straight, many, be- 1933J PINKERTON — FERNS OF MISSOURI 53 coming confluent with age, mostly solitary, occurring also on the slender prolongation. July to October. Distribution: Massachusetts to Alabama, westward to Missouri; recorded from Georgia and Arkansas. On La Motte sandstone only, in shaded crevices of cliffs. Specimens examined: Trelease, Engelmann, Russell, Pinkerton. 5. Asplenium platyneuron 2 (L.) Oakes in D. C. Eaton, Fern* N. Am. 1: 24. 1879. Gray, p. 39; Britton & Brown, ed. 1, 1 23, ed. 2, 1 : 27; Eaton, pi. 4, fig- *• Ebony spleenwort. Asplenium ebeneum Ait. Hort. Kew. 3: 462. 1789. Fronds 4r-20 inches high, fertile ones tall and upright, sterih ones short and spreading; stipes and rachises reddish brown shining, rather thick ; blades linear-oblanceolate, tapering at base pinnae 20-40 inches alternate, sessile, auricled on upper or both sides of base, and more or less overlapping rachis; sori 8-15 in number, nearer midveins than margins, becoming confluent. July to September. Distribution: Vermont to Alabama, westward to Texas and Oklahoma; recorded from Ontario and Colorado. Rocky open woods, preferring alkaline soil. Specimens examined: Davis, Kellogg, Engelmann, Eggert, Trelease, Bush, Palmer, Daniels, Pinkerton, Woodson, Russell, Mann, Dewart, Emig, Krause, Meek. Asplenium resiliens Kunze, Linnaea 18: 331. 1844. Gray ; Britton & Brown, ed. 1, 1 : 23, ed. 2, 1 : 27; Eaton, pi. 86, figs & Asplenium parvulum Mart. & Gal. in M6m. Acad. Brux. 15: 60. 1842. Rootstocks with black scales; fronds 4-12.5 inches long; stipes and rachises black and shining, slender; blades normally linear-oblanceolate, pinnate; pinnae 4-12 mm. long, mostly opposite, nearly sessile, upper edges auricled and on lower pinnae both edges auricled, deflexed; blades widest in middle; margins mostly entire or slightly crenate, tendency to incurve slightly; fruit dots nearer outer margins than midribs, nearly parallel to the midribs, oblong, few, sometimes becoming confluent. June to October. J Var. incisum (E. C. Howe) Robinson in Rhodora 10: 29. 1908, has very brittle stipes and the pinnae deeply pinnatifid. This appears to be merely an ecological variation. [Vol. 20 54 ANNALS OF THE MISSOURI BOTANICAL GARDEN Distribution: Massachusetts to Florida, westward to New Mexico. Limestone cliffs. Specimens examined: Trelease, Bush, Palmer, Russell, Pinkerton, Drouet. 7. Asplenium Trichomanes L. Sp. PI. 2: 1080. 1753. Gray, p. 30; Britton & Brown, ed. 1, 1: 24, ed. 2, 1: 28; Eaton, pi 36, fig. 1. Maidenhair spleenwort. Rootstocks nearly erect, inconspicuously chaffy with narrow black scales; fronds 3-6 inches high; stipes slender, densely tufted, brownish-purple, polished, rachis similar to tip; blades 7 mm. long, herbaceous, mos the opposite, roundish, crenate margins, obliquely wedge-trun at base, attached by narrow points; sori medial or nearer midveins than margins, 3-6 pairs on outer sides of veins, be- coming confluent; indusia membranaceous. July to September. Distribution: Ontario to Alabama, westward to British Columbia and California; widely distributed but local. Sandstone rocks where plenty of water is available. Specimens examined: Kellogg, Letterman, Eggert, Engelmann, Palmer, Russell, Pinkerton, Trelease, Morrison, liickett. Mackenzie. Blankenshi n . ATHYRIUM 1. Fronds simply pinnate /. A. angustifolium 1. Fronds more than pinnate 2 2. Fronds deeply bipinnatifid, margins lightly serrate-crenate 2. A. acroslichoides 2. Fronds usually tripinnatifid, margins deeply and irregularly incised 3 3. Rhizomes creeping, not densely covered with persistent leaf-bases; fronds widest near base; indusia with glandular cilia; spores nigrescent, wrinkled. 3. A. asplenioides 3. Rhizomes horizontal, completely concealed by thick fleshy bases of old fronds; fronds widest near middle; indusia toothed or short-ciliate, never glandular; spores yellow, slightly papillate J,.. A. angustum 1. Athyrium angustifolium (Michx.) Milde in Bot. Zeit. 48: 376. 1886. Gray, p. 3D; Britton & Brown, ed. 1, 1 : 24, ed. 2, 1 : 28; Eaton, pi 56, fig. 1. Narrow-leaved spleenwort. Asplenium pyenocarpon Spreng. Anleit. 3: 112. 1804. Asplenium angustifolium Michx. Fl. Bor. Am. 2: 265. 1803. Athyrium pyenocarpon Tidestrom, Elys. Marianum, p. 36. 1906. Rootstocks stout, creeping, with many long, branched rootlets; stipes green except for brown base; fronds 1-2.5 feet long, mem- branaceous, herbaceous, pinnate; pinnae 2-5 inches long, 20-30 pairs, short-stalked, linear-oblong, attenuate, margins slightly 1933] PINKERTON — FERNS OF MISSOURI 55 wavy; fertile pinnae near top, narrower and shorter; sori 20-30 pairs, linear, slightly curved, lying along outer of bifurcated veins; indusia firm, convex, concealed by strongly confluent sori at maturity. August and September. Distribution: Quebec to Georgia, westward to Michigan and Missouri; recorded from Kansas and Minnesota. Moist woods and shaded ravines, reported occasion- ally on sandy soil. Specimens examined: Davis, Kellogg, Palmer, Eggert, Trelease, Bush, Pinkerton, Daniels, Einig, Glatfelter. 2. Athyrium acrostichoides (Sw.) Diels in Engl. & Prantl, Nat. Pfl. V : 223. 1899. Gray, p. 39 ; Britton & Brown, ed. 1, 1 : 26, ed. 2, 1 : 30; Eaton, pi. 50. Silvery spleenwort. Asplenium acrostichoides Sw. in Schrad. Jour, fur die Bot. 2: (1800): 54. 1801. Asplenium thelypteroides Michx. Fl. Bor. Am. 2: 265. 1803. Athyrium thelypteroides (Michx.) Desv. in Mem. Soc. Linn. Paris [Prodr. p. 266] 6: 266. 1827. Rootstocks creeping, horizontal; stipes 8-16 inches long, straw-colored, herbaceous, with a few scales on lower portion; blades lanceolate to ovate-oblong, 1-3 feet long and 6-12 inches broad, narrowed to base, deeply bipinnatifid ; ultimate segments distinct, obtuse; margins slightly serrate-crenate ; sori 3-6 pairs per segment, arranged more or less evenly along lateral veins, mostly straight, oblong, some double. August to October. Distribution: New Brunswick to Georgia, westward to Missouri; recorded north- ward to Minnesota. Rich moist woods, or moist sandy soil. Specimens examined: Palmer, Eggert, Davis, Letterman, Broadhead. 3. Athyrium asplenioides 3 (Michx.) Desv. in Mem. Soc. Linn. Paris [Prodr. p. 266] 6: 266. 1827. Gray, p. 40; Britton & Brown, ed. 1, 1: 26, ed. 2, 1: 30; Butters in Rhodora 19: 169. 1917. Asplenium Athyrium Spreng. Anleit. 3: 113. 1804. Athyrium Filix-foemina (L.) Roth in Romer's Arch. f. Bot. 2 1 : 106. 1799. Rhizomes horizontally creeping, partially covered by short persistent leaf bases, the whole structure 1-1.5 cm. in diameter, * This species and A. angustum are very difficult to distinguish. It is often necessary to have the whole plant, fruiting and not too mature, to be absolutely certain. I have taken the character of the spore as my ultimate criterion. [Vol. 20 56 ANNALS OF THE MISSOURI BOTANICAL GARDEN with conspicuous projections of new growths before fronds of the current season; stipes long, about equal to the deltoid lanceolate fronds; young growths covered with scales soon deciduous, these small and light tan-colored, cell walls thin and inconspicuous; fronds bipinnate to tripinnatifid, second pair of pinnae commonly the longest, the lowest only slightly shorter; pinnae narrower at base; pinnules variously incised but apex more or less obtuse due to the venation, since two veins usually end on the same level; sori longer and narrower than in A. angustum, on the anterior side of anterior vein of each lobe of pinnule, and some- times on the lower veins, of typical athyroid type, rarely double ; young indusia ciliate with multicellular glandular-tipped hairs nearly disappearing at maturity, leaving quite even margins; sporangia stalk frequently supplied with a glandular hair; spores furnished with a somewhat nigrescent, wrinkled exospore. July to October. Distribution : Massachusetts to Florida, westward to Missouri and Texas. Shaded rich woods or cliffs, sandy soil. Specimens examined: Kellogg, Trelease, Palmer, Bush, Soulard, Blankenship. 4. Athyrium angustum 4 (Willd.) Presl in Rel. Haenk. 1: 39. 1825. Gray, p. 40; Britton & Brown, ed. 1, 1: 26, ed. 2, 1: 30; Butters in Rhodora 19: 169. 1917. Asplenium Michauxii Spreng. Syst. 4: 88. 1827. Asplenium Filix-foemina (L.) Roth in Romer's Arch. f. Bot. 2»:106. 1799. Rhizomes horizontal and somewhat oblique, condensed and completely covered by thick fleshy bases, 2-5 cm. in diameter; stipes up to half as long as the frond, a moderate number of scales persistent, 1 X 1.5 mm., dark opaque with thick darker cell walls and narrow cells; fronds bi-tripinnatifid, middle of fronds widest and lower pinnae much shorter and often deflexed, often large forms and polymorphic; pinnae not narrower at base; pinnules and segments with acute apices, due to one vein ending considerably beyond any of its neighbors; fertile fronds consider- 4 Butlers describes two varieties, namely: vur. elatius and var. rubellum, distin- guished chiefly by the dimorphism and thicker texture of the former (a sun form) and the nionornorphism and thinner texture of the latter (a shade form). He claims that the latter has a more northerly distribution; but in Missouri the two seem to be purely ecological variations and scarcely worth varietal rank. 1933) PINKERTON FERNS OF MISSOURI 57 ably narrower and more acute than sterile ones; son short and wide, of typical athyroid type; indusia never glandular but persistently short-ciliate ; sporangia stalk rarely bearing glandular hair ; spores bright yellow and slightly papillate, no exospore and not wrinkled. June to October. Distribution: Labrador to Pennsylvania, westward to South Dakota and Missouri. Rich woods. Specimens examined: Bush. Davis. Palmer. Enaelmann. Kelloaa. EaaerL Russell. CAMPTOSORUS 1. Camptosorus rhizophyllus (L.) Link, Hort. Berol. 2: 69. 1833. Gray, p. 40; Britton & Brown, ed. 1, 1: 21, ed. 2, 1: 26; Eaton, pi. 8, fig. 1. Walking fern. Rootstocks short, creeping; stipes grouped in tufts, spreading, green, fleshy, 1-6 inches long; blades evergreen, subcoriaceous, 4-12 inches long, base auriculate, cordate or hastate, apex attenuate and filiform, rooting at tips (or from auricles); sori numerous, straight or slightly curved, single and on inside of and parallel to veins, near midribs, on both sides of outer veins and becoming confluent at exterior tips; indusia membranaceous. All year. Distribution: Ontario to Alabama, westward to Minnesota and Oklahoma. Lime- stone rocks, usually associated with Entodon (a moss). Specimens examined: Bush, Davis, Engelmann, Trelease, Kellogg, Daniels, Palmer, Pinkerlon, Dewart, Russell, Rickett, Weller. CHEILANTHES 1. Fronds relatively smooth 1. C. alabamensis 1. Fronds hairy or tomentose 2 2. Plants small, 2-6 inches tall, matted; mature stipes nearly glabrous. .2. C. Feei 2. Plants taller, 4—20 inches, not matted; stipes hirsute 3. C. lanosa 1. Cheilanthes alabamensis (Buckl.) Kunze in Linnaea 20: 4. 1847. Gray, p. 36; Britton & Brown, ed. 1, 1: 30, ed. 2, 1: 34; Eaton, pi. 57, fig. 7. Rootstocks creeping, short, slender, with dark ferruginous scales; stipes black, wiry, slightly villous at base due to fine rusty scales 3-7 mm. long; blades scabrous to smooth, 2-10 inches long, lanceolate, bipinnate; pinnae acuminate, lower ones shorter than those above; pinnules often auriculate; indusia broad, pale but firm, frequently broken by incision of pinnules. August to October. [Vol. 20 58 ANNALS OF THE MISSOURI BOTANICAL GARDEN Distribution: Florida, westward to Missouri and Arizona. Limestone cliffs. Specimens examined: Kellogg, Palmer, Bmh. 2. Cheilanthes Feei Moore, Ind. Fil, Gen. 38. 1857. Gray, p. 36; Britton & Brown, ed. 2, 1: 34; Eaton, pi. 6, jig. 1. Cheilanthes lanuginosa Nutt. in Hk. Sp. Fil. 2: 99. 1852. Rootstocks short, clothed with narrow scales of black centers and thin brown edges; fronds 2-6 inches tall; stipes densely tufted, black or brown, originally woolly, becoming glabrous when mature; blades bi-tripinnate, slightly lomentose above and woolly below; pinnules divided into minute rounded segments, densely crowded; indusia herbaceous, continuous. July to Oc- tober. Distribution: Wisconsin to Texas, westward to Nevada and Arizona. Limestone cliffs in dry and exposed localities. Specimens examined: Davis, Kellogg, Engdmann, TnUase, Bush, Palmer, Daniels, Pinkerton, Morrison, Uphof. 3. Cheilanthes lanosa (Michx.) Watt in Jour. Bot. 12: 48. 1874. Gray, p. 30; Britton & Brown, ed. 1, 1: 31, ed. 2, 1: 34; Eaton, pi 2, fig. 2. Cheilanthes vestita Sw. Syn. Fil. 128. 1806. Rootstocks short, creeping, with pale brown scales; fronds 4-10 inches long; stipes wiry, dark brown, hirsute; blades her- baceous, bipinnate; pinnae hirsute and somewhat glandular; in- dusia inconspicuous, discontinuous. July to September. Distribution: New York to Alabama, westward to Oklahoma; recorded from Connecticut and Texas. Sandstone rocks, usually in dry and exposed places. Specimens examined: Kellogg, Eggert, Ldterman, Palmer, Broadhead, Pinkerton, Midler, (ilatfeltcr, Mackenzie, Swallow, Emzg, Blankenship, Link. CYSTOPTERIS 1. Pinnae* short-stalked on rachises; pinnules at least narrowed at points of attachment; segments ovate, acute, usually variously incised; indusia truncate 1. C. fragilis 1. Pinnae sessile on rachises; pinnules oblong, obtuse, regularly toothed; indusia round or pointed at apex 2 2. Fronds long-attenuate 2. C. bulbifera 2. Fronds not long-attenuate 3. C. bulbifera var. horizontalis 1. Cystopteris fragilis 5 (L.) Bernh. in Schrad. Neues Jour. 6 An exceedingly variable species. Hybridization with C. bulbifera might account for some of the aberration. Forms bearing bulblets have been included under this group, as that character does not seem to be a constant feature for C. bulbifera alone. 1933] PINKERTON — FERNS OP MISSOURI 59 Bot. I 2 : 27. 1806. Gray, p. 43; Britton & Brown, ed. 1, 1: 13, ed. 2, 1 : 15; Eaton, pi 53, fig. 1. Filixfragilis Underw. Nat. Ferns, ed. 6, 119. 1900. Rootstocks elongated, often 4-5 inches long, or shorter and condensed, slender, but covered with persistent leaf-bases, chaffy at apex, scales delicate, ovate, acuminate, ferruginous; stipes in a dense cluster, slender, brittle, 4-6 inches long; blades 6-8 inches long and about half as wide, thin and membranaceous, ovate-lanceolate; basal pinnae commonly narrower than the second and third pairs, apparently bipinnate but segments usually connected by narrowly winged midribs, segments round- ish-oval to ovate to rhomboid-ovate to ovate-lanceolate, toothed, dentate or irregularly laciniate; veinlets pinnately arranged on mid veins, lower ones forked; sori small, roundish, seated on middle of veins nearest midrib ; indusia delicate, rounded, ovate, or occasionally with narrow beak-like points, concealed by mature sporangia; lower pinnae often sterile. May to July. Distribution: Cosmopolitan. Rocky soil, moist woods, pref erring alkaline soil. Specimens examined: Davis, Engelmann, Trelease, Bush, Palmer, Pinkerton, Eggert, Kellogg, Woodson, Harrison, Mann, Daniels, Blankenshiv. Duncan. Williams. 2. Cystopteris bulbifera (L.) Bernh. in Schrad. Neues Jour. Bot. I 2 : 10. 1806. Gray, p. 43; Britton & Brown, ed. 1, 1: 12, ed. 2, 1 : 15; Eaton, pi. 53, fig. 13. Filix bulbifera Underw. Nat. Ferns, ed. 6, 119. 1900. Rootstocks seldom over one inch long, chaffy at apex and covered with persistent leaf-bases; stipes slender, rather brittle, 6-10 inches long; blades submembranaceous but of a brittle rigidity, triangular-attenuate, 1-4 feet long, 3-5 inches broad at base; pinnae attenuated upward, bipinnate at base, upper pinnules attached by winged rachises; pinnae numerous (up to 40 pairs), oblong; pinnules oblong, obtuse, pinnately lobed; sori numerous, all pinnae fertile, arranged in rows along each side of midveins of pinnules, placed on the lowest superior veinlet of each group near its middle and so near the midvein; indusia truncate and fragile, covered by mature sporangia; bulblets some- times present on the under side of the frond attached near the base of or on the pinnae. July to August. Distribution: Newfoundland to Georgia, westward to Michigan and Arkansas: (Vol. 20 60 ANNALS OF THE MISSOURI BOTANICAL GARDEN recorded from Utah and Arizona. Rocky soil, preferring limestone, in moist shady situations. Specimens examined: Bush, Davis, Kellogg, Trelease, Palmer, Pinkerton, Daniels, Letterma n. 3. CvstODteris bulbifera var. horizontalis Lawson in Bot. Soc. Edinb. Trans. 8:40. 1866. PL 5, fig Fronds triangular-lanceolate, broad at base, not more than three or four times longer than broad; pinnae horizontal; lowest Dinnules often ouite broad with irregularly cut lobes and bearing us medium-sized sori along the tripinnate, or pinnules irregula lobed, m lanceolate. May to October. Distribution: southern Missouri and northern Arkansas. Damp limestone bluffs. Specimens examined: Ilasco, Ralls Co., Davis 2668; Sulphur Springs, Jefferson Co., coll. of Oct. 23, 1898, Trelease; Terre Bleue Cr., Ste. Genevieve Co., coll. of Aug. 29, 30, 1898, Trdaise; Lesterville, Reynolds Co., coll. of June 5, 1929, Kellogg; Tecumseh, Ozark Co., Palmer S2896. DENNSTAEDTIA 1. Dennstaedtia punctilobula (Michx.) Moore, Ind.Fil. Gen. 97. 1857. Gray, p. 45; Britton & Brown, ed. 1, 1: 12, ed. 2, 1: 14; Eaton, pi. 44- Dicksonia pilosiuscula Willd. Enum. 1076. 1809. Dicksonia punctilobula Hk. Sp. Fil. 1 : 79. 1846. Rootstocks slender, scaleless but finely hairy at tips, irregularly branching with many long slender root- lets; stipes rather stout, light brown, chaffless, slightly puberulent; blades 1-3 feet long, ovate-lanceolate, acuminate, delicately herbaceous, hairy and minutely glandular, tripinnatifid ; pinnae numerous, lanceolate, pointed, second pair a little longer than first; pinnules adnate to rachis and usually decurrent on rhomboid-ovate, pinnatifid into oblong and obtuse cut-toothed lobes; sori minute, on upper margins of the lobes of the pinnules; indusia cup-like, delicate. August. Distribution: New Brunswick to Georgia, westward to Illinois and Missouri. Civvices of La Motte sandstone, rare. Specimens examined: Trelease, Russell, Kggert, Pinkerton. NOTHOLAENA 1. Notholaena dealbata (Pursh) Kunze in Amer. Jour. Sci. II. 6: 82. 1S48, as Nothochlaena. Gray, p. 35; Britton & Brown, ed. 1, 1 : 32, ed. 2, 1 : 35; Eaton, pi. 9, fig. 2. 1933] PINKERTON — FERNS OF MISSOURI 61 Cheilanlhes dealbala Pursh, Fl. Am. Sept. 2: 671. 1814. Nothochlaena pulchella Kunze in Bot. Zeit. 1 : 633. 1843. Pellaea dealbata (Pursh) Prantl in Engler's Bot. Jahrb. 3: 417. 1882. Notholaena nivea var. dealbata Davenp. in Cat. Davenp. Herb. Suppl. 44. 1883. Rootstocks short, creeping, with narrow brown chaffy scales; stipes 1-4 inches long, tufted, wiry, slender, copper-brown, as are the rachises; blades 2-4 inches long, broadly deltoid-ovate, 4-5- pinnate, all but ultimate segments alternate, those sometimes opposite ; segments obovate-oval and entire or several-lobed, 1-2 mm. broad, upper surfaces pale green, coriaceous, lower white and powdery, giving a silvery appearance; sporangia seated on upper portions of the veins ; no indusia, but a protection afforded by the slightly turned-back margins. June to September. Distribution: Missouri and Kansas to central Texas; recorded from Nebraska. Dry calcareous rocks. Specimens examined: Daniels, Bush, Palmer, Blankenship. ONOCLEA 1. Onoclea sensibilis L. Sp. PL 2: 1062. 1753. Gray, p. 45; Britton & Brown, ed. 1, 1: 9, ed. 2, 1: 11; Eaton, pi. 72, fig. 1. Sensitive fern. Rootstocks slender, creeping, rooting freely and often forking; stipes coarse, straw-like, hollow, flattened, light brown when dry; sterile blades triangular or triovate, midribs winged, widening toward the apex, sinuses rounded; lowest segments broadly lanceolate, herbaceous, sensitive to frost; veins conspicuous, re- ticulate; margins variously rounded, lobed, toothed, serrate or fronds contracted: each segment a pouch filled with several sporangia; delicate hood-like indusia. August to November. Newf Swamps or damp rich soil. Palmer. Morrison Blankenship, Mackenzie, Williams. PELLAEA 1. Pinnae dichotomously branched at apex S. P. atropurpurea var. cristata 1. Pinnae not branched at apex 2 [Vol. 20 62 ANNALS OF THE MISSOUKI HOTANICAL GARDEN 2. Stipes smooth, reddish-brown; pinnae usually membranaceous, pale green, short, more or less roundel at both corners and tending to divide at bases into two or more parts /. p. glabella 2. Stipes scabrous, dark purplish-black ; pinnae coriaceous, blue-green, elongate 1 seldom redivided 2. P. atropurpurea an< 1. Pellaea glabella Mett., Kuhn in Linnaea 36: 87. 1869. Gray, p. 37; Britton & Brown, ed. 2, 1: 33. Pellaea atropur purea var. Buxkii Mackenzie, Flora Jackson ( iounty, Mo., p. 5. 1902. Stipes and rachises brownish-red, smooth or but slightly hairy; fronds simply pinnate above, the lower ternate or rarely quinate (the entire pinnule seems to be breaking at the base— a possible tendency loward compound pinnules); pinnae membranaceous to but April to October Distribution: Ontario and Vermont to Pennsylvania, westward to South Dakota and northern Arkansas. Exposed high places on limestone cliffs. Specimens examined: Davis, Kggeri, Kellogg, Bush, Palmer, Pinkcrton, Trelease. 2. Pellaea atropurpurea (L.) Link, Fil. Sp. in Hort. Berol. 59. 1841. Cray, p. 37; Britton & Brown, ed. 1, 1: 29, ed. 2, 1: 33; Eaton, pi. 64, fig. 4> Rootstock short, densely covered with rusty scales about 2 mm. in length ; fronds 4-12 inches long, pinnate or below bipinnate, coriaceous; fertile segments linear, more or less pointed at apex; sterile segments approaching oval, shortly stalked; veins obscure; continuous indusia of reflexed margins. June to September. Distribution: Connecticut to Florida, westward to South Dakota and Texas. Near small limestone rocks or on top of cliffs where there is loose soil. Specimens examined: Davit, Engelmann, Trelease, Broad head, Bush, Palmer, Pinkcrton, Kellogg, Daniels. 3. Pellaea atropurpurea var. cristata Trel. in Rept Card. 12:77. 1901. Cray, p. 37. Pinnae dichotomously forked. Distribution: known only from Eureka, Missouri. Limestone. Specimens examined: Eureka, 1899, Pauls. POLYPODIUM 1. Blades smooth, green /. />. virginianum 1. Blades densely scaly, grayish 2. P. poly podioides 1933] PINKERTON — FERNS OF MISSOURI 63 1. Polypodium virginianum L. Sp. PI. 2: 1085. 1753. Gray, p. 34; Britton & Brown, ed. 1, 1: 32, ed. 2, 1: 36; Eaton, pi. 81, fig. 1. Common polypody. Polypodium vulgare L. Sp. PL 2: 1085. 1753. Rootstocks close to surface of soil, covered with chaffy, red- brown scales ; stipes smooth, herbaceous, light green ; blades ovate- oblong or narrowly oblong, subcoriaceous or chartaceous, ever- green, simple, deeply pinnatifid, smooth ; segments linear-oblong, obtuse or slightly acute, crenulate and serrate, sinuses rounded, alternate, margins obscurely dentate; sori large, naked. July. Distribution: Newfoundland to Florida, westward to Minnesota and Arkansas. On sandstone or sandy soil. Specimens examined: Eggert, Russell, Engelmann, Trelease, Pinkerton, Letterman, Greene. 2. Polypodium polypodioides (L.) Hitchc. in Rept. Mo. Bot. Gard. 4: 156. 1893. Gray, p. 34; Britton & Brown, ed. 1, 1 : 33, ed. 2, 1 : 36; Eaton, pi. 26, fig. 2. Gray polypody. Polypodium incanum Sw. Fl. Ind. Occ. 3: 1645. 1806. Rootstocks woody, covered with small dark brown scales; stipes 1-4 inches long, slender, bearing peltate ovate scales with dark brown centers; blades oblong-lanceolate, pinnate; segments ob- long, obtuse, entire, sessile, separated by rounded sinuses, alter- nate, upper surfaces smooth or with few scales, lower densely scaly ; fruit dots small and naked. July to September. Distribution : Massachusetts to Florida, westward to Missouri and Texas. Usually Taxodium rocks. Mac kenzie, Meek. POLYSTICHUM 1. Margins serrulate; fertile pinnae contracted; sori confluent. ...UP. acroslichoides 1. Margins deeply toothed or pinnatifid; fertile pinnae scarcely contracted; sori not confluent and appearing only on tips of lower pinnae 2. P. acrostichoides var. incisum 1. Polystichum acrostichoides (Michx.) Schott, Gen. Fil. 17. 1834. Gray, p. 40; Britton & Brown, ed. 1, 1: 14, ed. 2, 1: 16; Eaton, pi. 84- Christmas fern. Aspidium acrostichoides Sw. Syn. Fil. 44. 1806. Dryopteris acrostichoides Kuntze, Rev. Gen. PL 2:812. 1891 . 64 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 20 Root-stocks stout, creeping, with persistent leaf-bases; stipes densely tufted, with chaff of large golden-brown scales, 5-7 mm. wide; blades lanceolate, pinnate, 1-2 feet long, 3-5 inches wide, rigid, evergreen, subcoriaceous ; pinnae numerous, 1-3 inches long, oblong-lanceolate, short-stalked, upwardly falcate or lowest slightly dellexed, apex acutish, upper sides auriculate; margins serrulate to incised with incurved bristle-pointed teeth; veins free, branching three to four times; upper pinnae of fertile fronds more or less contracted and heavily soriferous; sori terminal on lower veinlets in 2-4 rows, becoming confluent with age; indusia round, indurated, not glandular, persistent. July to August. Distribution: Maine to Florida, westward to Michigan and Texas; recorded from Nova Scotia. Shady hillsides of ravines, in rich soil which is interspersed with rocks; common. Specimens examined: Bush, Davis, Eggert, Trelease, Palmer, Pinkerton, Emig, Kellogg, Darnels, Thomas, Mackenzie. 2. Polystichum acrostichoides var. incisum Gray, Man. Bot. ed. 1, 032. 1848. Gray, p. 40; Britton & Brown, ed. 1, 1: 14, ed. 2, 1: 10. Polystichum acrostichoides var. Schweinitzii (Beck) Small in Bull. Torr. Bot. Club 20: 404. 1893. Aspidium schweinitzii Beck, Bot. North. & Mid. States, ed. 1 , 449. 1833. Aspidium acrostichoides var. incisum D. C. Eaton, Ferns N. Am. 1:258. 1879. Segments few and distant, large, irregularly incised; upper pinnae covered by confluent sori, lower ones fertile at tips only, sori large; veins numerous, frequently overlapping to form ir- regular areoles. Distribution: This seems to be nearly as common as the type in southern Missouri and northern Arkansas; but since there appears to be a series of intermediate forms between the two I have not attempted to separate the individual ranges. PTERETIS 1. Pteretis nodulosa (Michx.) Nieuwl. in Am. Midi. Nat. 4: 334. 1910. Gray, p. 45; Britton & Brown, ed. 1, 1:9, ed. 2, 1 : 11; Eaton, pi. 73. Ostrich fern. (kmuhda Struthiopteris L. Sp. PI. 2: 1000. 1753. Onoclca Struthiopteris Hoffm. Deutsch. Fl. 2:11. 1795. 1933] PINKERTON — FERNS OF MISSOURI 65 Struthiopteris germanica Willd. Enum. 1071. 1809. Malteuccia Struthiopteris (L.) Todaro in Syn. PL Acot. Vase. Sicilia, p. 30. 1866. Matteuccia nodulosa (Michx.) Fernald in Rhodora 17: 164. 1915. Rootstocks stout and ascending, with slender underground stolons; sterile blades 2-7 feet high, 6-15 inches broad, short- stalked, broadly oblanceolate, abruptly short-acuminate, gradual- ly narrowed below middle, lower pinnae reduced ; pinnae narrow, deeply pinnatifid; segments oblong, obtuse, entire; veins simple, fertile blades 1-7.5 feet high, with pod-like brown pinnae, in- cluded sori confluent. July. Distribution: Newfoundland to New York, westward to Michigan and Missouri; recorded from Virginia. Alluvial soil. Specimens examined: Livonia, Putnam Co., Bush 7780, 7780 A, 7780B. PTERIDIUM 1. Pinnules much elongated, seldom redivided . . .2. P. latiusculum var. pseudocaudatum 1. Pinnules oblong-lanceolate, at least a few pinnatifid 1. P. latiusculum 1. Pteridium latiusculum (Desv.) Hieron. Wissenschaftl. Ergeb. d. Schwed. Rhodesia-Kongo-Exp. 1911-12, Heft 1, p. 7. 1914; Maxon in Am. Fern Jour. 9: 43. 1919. Gray, p. 36; Britton & Brown, ed. 1, 1 : 28, ed. 2, 1 : 32; Eaton, pi. 85. Pteris aquilina L. Sp. PL 2: 1075. 1753. Pteridium aquilinum Kuhn in Decken, Reisen in Ost-Afrika 3 3 : 11. 1879. Rootstocks cord-like, creeping, blackish, deeply buried; stipes solitary, erect, naked, 1-3 feet high, swollen at base, brownish; blades 2-4 feet long, 1-3 feet broad, triangular-ovate, rigidly subcoriaceous, subternate; the long-stalked basal pinnae and middle ones bipinnate, those above lobed or simple; segments oblong-lanceolate, under surfaces glabrous or pubescent; veins close-packed, free; indusia continuous around the edge of the pinnules, double. July to September. Distribution: Cosmopolitan, open woods, preferring sandy soil. Specimens examined : Kellogg, Eggert, Palmer, Bush. 2. Pteridium latiusculum var. pseudocaudatum Maxon in Am. Fern Jour. 9 : 44. 1919. Gray, p. 36; Britton & Brown, ed. 2, 1:32. [Vol. 20 66 ANNALS OF THE MISSOURI BOTANICAL GARDEN The variety occurs in this region but does not seem to have a distribution different from that of the species. It is distinguished by its very narrow elongated pinnules. TIIELYPTERIS Dryopteris Adans. Polystichum Roth Aspidium Swartz Nephr odium Richard Lastrea Bory Phegopteris (Presl) Fee 1. Indusia absent 5. T. hexagonopiera 1. Indusia present 2 2. Blades bipmnatifid or bipinnate; segments not spinulose 3 2. Blades tripinnatifid or tripinnate; segments spinulose 4 3. Sori medial, small; blades membranaceous, not evergreen 1. T. palustris var. pubescens 3. Sori near margins, large; blades subcoriaceous, evergreen 2. T. marginalia 4. Indusiu glandless; pinnae decidedly oblique to rachises, scales of stipe pale brown s. t. sjnnul 4. Indusia glandular; pinnae more or less at right angles to rachises; scales of osa stipe brown with a dark center 4. T. sidnulosa var. intermedia 1. Thelypteris palustris var. pubescens 6 (Lawson) Fernald in Rhodora 31: 34. 1929. Gray, p. 41; Britton & Brown, ed. 1, 1 : 15, ed. 2, 1 : 18; Eaton, pi. SO. Marsh shield-fern. Thelypteris Thelypteris Nieuwl. in Am. Midi. Nat. 1 : 226. 1910. Root stocks slender, creeping, blackish, nearly naked; stipes as long or longer than the blades, blackish at base, sparingly chaffy; blades oblong-lanceolate, pinnate, 1-3 feet long, 4-6 inches wide, membranaceous, scarcely narrowed at base, short- acuniiiuite; pinnae 20-30 pairs, alternate, short -stalked, approxi- mately at right angles to stalks, linear-lanceolate, base, deeply pinnatifid; segments oblong-obtuse, m broadest veins pinnate, usually once-forked; fertile fronds usually on longer stalks and of narrower segments than the former; sori nearly medial, crowded; indusia glabrous. August. 6 Eaton mentions an unusual form — "the lower two or three pairs are usually but little shorter than those above them; but fronds are occasionally found in which they are conspicuously reduced." Such a form is one from Iron Lake, Iron Co., Kellogg \6SL 1933) PINKERTON — FERNS OF MISSOURI 67 Distribution : New Brunswick to Florida, westward to Texas. Wet Specimens examined: Eggert, Kellogg, Bush, Pinkerton. 2. Thelypteris marginalis (L.) Nieuwl. in Am. Midi. Nat. 1: 226. 1910. Gray, p. 42; Britton & Brown, ed. 1, 1: 17, ed. 2, 1 : 20; Eaton, pi. 55. Evergreen wood fern. Rootstocks stout, ascending, covered with long chaffy brown scales; stipes several inches to a foot long, light tan, somewhat chaffy; blades 6-30 inches long, evergreen, subcoriaceous, ovate- lanceolate, scarcely narrowed at base, deeply bipinnatifid ; pinnae numerous, practically sessile, lanceolate, acuminate, slightly broader above the base ; pinnules adnate to narrowly winged sec- ondary rachis, oblong to oblong-lanceolate, faintly crenately- toothed ; veins free, forked or pinnately branched ; sori large, near margins of segments; indusia hard, orbicular-reniform, glabrous, dark brown. July to August. Distribution : Nova Scotia to Georgia, westward to Kansas and Oklahoma. Sand- stone ledges where it is moderately moist. Specimens examined: Letterman, Engelmann, Eggert, Kellogg, Palmer, Pinkerton, Midler, Rickelt, Mackenzie, Broadhead, IAnk, Blankenship, Trelease. 3. Thelypteris spinulosa (Retz.) Nieuwl. in Am. Midi. Nat. 1 : 226. 1910. Gray, p. 43; Britton & Brown, ed. 1, 1: 18, ed. 2, 1:21; Eaton, pi. 68. Spinulose shield-fern. Rootstocks stout, creeping, chaffy; stipes 4-14 inches long, chaffy; blades 0.5-3 feet long, ovate-lanceolate to oblong, acumi- nate, bi-tripinnate, firmly membranaceous ; primary pinnae short- stalked, lower pairs triangular-ovate to triangular-lanceolate, re- maining pinnae gradually narrower in outline; secondary rachises narrowly wing-margined; pinnules oblong, subacute, incised with spinulose-serrate lobes ; sori small, sub-marginal, terminal on vein- lets; indusia flat, round-reniform, glandless. Distribution: Labrador to Virginia, westward to Idaho. Moist woods, alluvial soil. Specimens examined: Neeleyville, Butler Co., coll. of Oct. 30, 1899, Russell. 4. Thelypteris spinulosa var. intermedia (Retz.) Nieuwl. in Am. Midi. Nat. 2: 278. 1912. Gray, p. 43; Britton & Brown, ed. 2, 1 : 22. Similar to the species except for the glandular indusia and right-angled relation of pinnae to rachis. [Vol. 20 68 ANNALS OF THE MISSOURI BOTANICAL GARDEN Distribution: In Missouri — more northern, sandy soil. Specimens examined: Pickle Springs, Ste. Genevieve Co., Pinkerton 1; and coll. of May 21, 1930, Kellogg. 5. Thelypteris hexagonoptera (Michx.) Weatherby in Rhodora 21: 179. 1919, Gray, p. 35; Britton & Brown, ed. 1, 1: 19, ed. 2, 1: 23; Eaton, pi. 65. Broad beech-fern, Phegopteris hexagonoptera (Michx.) F6e, Gen. Fil. 243. 1850-2. Rootstocks elongated, slender, creeping, chaffy with gold scales; stipes 8-20 inches long, slender, greenish, or straw-colored; blades triangular, 7-12 inches long, 7-15 inches wide, thinly herbaceous, deep green, slightly hairy or granular, bipinnatifid; pinnae adnate to winged rachis, the lowermost ones broadest and largely ovate to ovate-lanceolate, and others lanceolate; seg- ments usually bluntly acuminate and crenate; veins pinnate and free, branched or not; sori borne over whole frond, small, near end of veins and so near margins; no indusia. August. Distribution: New Brunswick to Delaware, westward to Oklahoma. Moist woods and ravines. Specimens examined: Davis, Eggert, Palmer, Kellogg, Bmh, Pinkerton, Rickett, Link. WOODSIA 1. Woodsia obtusa (Spreng.) Torr. Cat. PL in Geol. Rept. N. Y. 195. 1840. Gray, p. 44; Britton & Brown, ed. 1, 1: 11, ed. 2, 1: 14; Eaton, pi 71. Rootstocks short, creeping, chaffy with narrow light brown scales; stipes 2-6 inches long, green with darkish base in living plants and drying to a bright brownish straw color; fronds 8-15 inches long, broadly lanceolate, narrower at base than in middle, abruptly terminating at tip, membranaceo-herbaceous, minutely glandular, deep blue-green in color, nearly bipinnate; pinnae remote, short-stalked, obtuse, triangular, ovate to oblong, mostly opposite; segments oblong, obtuse, crenately toothed; sori sub- terminal on veins, nearer margin than mid veins; young indusia subglobose, splitting into several irregular lobes which extend out beyond the sporangia, difficult to detect. September. Distribution: Vermont to Alabama, westward to Wisconsin and Texas; recorded from Nova Scotia. Moist calcareous or acid soil. Specimens examined: Davis, Kellogg, Palmer, Pinkerton. 1933] PINKERTON — FERNS OF MISSOURI 69 WOODWARDIA 1. Woodwardia areolata (L.) Moore, Ind. Fil. Gen. 45. 1857. Gray, p. 38; Britton & Brown, ed. 1, 1: 20, ed. 2, 1: 25; Eaton, pi. 22, jig. 2. Lorinseria areolata (L.) Presl in Epim. Bot. 72. 1849. Acrostichwn areolatum L. Sp. PL 2: 1069. 1753. Woodwardia angustifolia J. E. Smith in Mem. Acad. Turin 5:411. 1793. Rootstocks creeping, several to 12 inches long, often branched, less than 1/4 inch thick, with some scales near apex; stipes dark at base, paler above, bearing a few scales, dimorphic; sterile fronds 9-10 inches long, oblong-ovate, pinnate, bright green above, paler below; rachis winged from tip to just below base of blades; sinuses rounded, segments acute, 1-4 inches long, 0.5-1 inch wide, finely serrate, membranaceous ; veins finely reticulated, with a longitudinal row of narrow areoles along each side of midribs and midveins, and several rows of hexagonal aeroles and free veins running outwards to serrated edges; fertile fronds taller, with a darker stalk ; segments and wing of rachis much narrowed ; one row of areoles on each side of midribs, each covered by a brown involucre attached to the outer enclosing veins and open along midrib; sporangia also from enclosing veinlets; sterile fronds appearing in May and fertile ones later. August to October. Distribution: Massachusetts to Florida, westward to Missouri and Texas; re- corded from Maine. Swamps and moist soil. Specimens examined: Poplar Bluff, Butler Co., July, 1898, Eby. Equisetaceae EQUISETUM 1. Stems annual; dimorphic i- E. arvense 1. Stems perennial; monomorphic 2 2. Sheaths cylindrical, green, turning gray, with black lines at bases and tops, short and undilated, splitting with age; ridges almost smooth; often very large plants #• %■ praealtum 2. Sheaths funnel-shaped, green with narrow black limbs, elongate, not splitting; ridges with one row of tubercules; medium-sized plants 8.E. laevigatum Equisetum arvense L. Sp. PL 2: 1061. 1753. Gray on & Brown, ed. 1, 1 : 36, ed. 2, 1 : 39. 70 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 20 Rootstock tuberiferous, felted with brown wool, extensively creeping; fronds dimorphic; sterile ones annual, prostrate or erect, green, rather slender, 12-24 inches high, 6-19 furrowed, with scattered stomata; sheaths whitish, tipped with about 12 acuminate, brown, separate teeth; branches whorled, simple or compound, not drooping, the 3-4 angled sheaths of branches consisting usually of 4 teeth, or often 3, rarely 5, long and acumin- ate; fertile fronds annual, appearing in early spring before sterile ones, usually unbranched, succulent, and withering after spores are ripe, 4-10 inches high, light brown, sheaths conspicuous, long, flaring and pointed, of 8-12 teeth; spikes not apiculate; variable. May, Distribution: Greenland to Alabama, westward to Alaska and California. Sandy soil, in waste places, along streams, etc. Specimens examined: Davis, Kellogg, Bush, Daniels, Paltrier, Letterman, Eggert, Trelease, Daniels, Mackenzie. 2. Equisetum praealtum 7 Raf. Fl. Ludovic. 13. 1817. Gray, p. 53; Brit ton & Brown, ed. 1, 1 : 38, ed. 2, 1 : 41. Equisetum hyemale var. robustum A. A. Eaton in Fern Bull. 11:74. 1903. Hippochaete prealta (Raf.) Farwell in Mem. N. Y. Bot. Gard. 6:467. 1916. Equisetum robustum A. Br., Engelm. in Amer. Jour. Sci. 46: 88. 1844. feet high, erect; stems rough drical Fronds perennial, evergreen, 3-11 20-48 ridges bearing silica in sin£ short-appressed, not dilated or only slightly when young, 8 at first green but soon turning black or gray with black bands above and below, splitting with age; sheath segments normally tri- carinate; teeth dark and caducous; cones pointed. May. 7 SchafTner (Am. Fern Jour. 13: 33-41. 1923), says: "Although perennial, E. praealtum usually bears cones on shoots of the season. Shoots sterile the first year may hear cones the second, both terminal and on lateral branches. Branching is rare the first season unless the shoot is injured, but the second year branching is common even on uninjured shoots. E. praealtum is an exceedingly variable species, some forms recognized probably being genetic and some ecological, but none of these forms passes out of the specific limits as usually drawn. Some are short and robust; souk tall and massive; some very slender." 8 Young shoots are often very difficult to distinguish from E. laevigatum. 1933] PINKERTON — FERNS OF MISSOURI 71 Distribution: Quebec to Georgia, westward to British Columbia and New Mexico. Wet sandy places. Specimens examined : Davis, Engelmann, Bush, Palmer, Pinkerton, Eggert, Trelease, Daniels, Throuse, Demetrio. 3. Equisetum laevigatum A. Br., Engelm. in Amer. Jour. Sci. 46: 87. 1844. Gray, p. 53; Britton & Brown, ed. 1, 1: 38, ed. 2, 1 : 42. Fronds perennial, erect, mostly simple, pale green, 1-5 feet high, 14-30 ridged, almost smooth, with stomata in two rows on each side of depressions; sheaths funnel-shaped, elongated, and green with usually a narrow black band at the top ; white-margined teeth soon deciduous; cones pointed. May to June. Distribution: New York to North Carolina, westward to Washington and Cali- fornia. Along streams, especially in sandy soil. Specimens examined : Engelmann, Bush, Daniels, Eggert, Palmer. OSMUNDACEAE OSMUNDA 1. Sterile fronds truly bipinnate; pinnules stalked and widely separated 1. O. regalis var. spectabilis 1. Sterile fronds bipinnatifid 2 2. Fronds dimorphic; sterile pinnae with tufts of brown hairs at base; apices of fronds and pinnae tapering; veins inconspicuous 2.0. dnnamomea 2. Fronds monomorphic but fertile part of frond is the middle several pairs of pinnae; no tufts of hair at base of pinnae; apices of fronds and pinnae abruptly narrowed and scarcely acute; veins dark-colored, conspicuous. 3. 0. Claytoniana 1. Osmunda regalis var. spectabilis Fernald in Rhodora 32: 72. 1930. Gray, p. 46; Britton & Brown, ed. 1,1:5, ed. 2, 1 : 7. Rootstocks stout, creeping, covered with persistent leaf-bases; fronds 2-6 feet high, stipes never chaffy, green or yellow, rounded on back, flattened on front; blades ovate-oblong, bipinnate; pinnae mostly opposite, ovate; pinnules unequal, 6-12 pairs plus the terminal one, subcoriaceous, short-petioled, distant, oval- oblong or oblong-lanceolate, ultimate ones often auriculate on lower side, margins crenulate-serrate, apex obtuse or subacute; apical pinnae fertile, bipinnate, ultimate divisions thread-like, containing no chlorophyll, entirely covered with sporangia. May to July. 72 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 20 Distribution: Newfoundland to Florida, westward to Saskatchewan and Missis- sippi. Lowlands, swamps, marshes, and wet woods. Specimens examined: Eggert, Palmer, Kellogg, Bush, Pinkerton, Engelmann, Trelease, Mackenzie. 2. Osmunda cinnamomea L. Sp. PL 2: 1066. 1753. Gray, p. 47; Britton & Brown, ed. 1, 1: 5, ed. 2, 1:7. Rool stocks creeping, massive, bearing circular clusters of sterile leaves with one or more fertile ones within; fronds dimor- phic, sterile ones 1 foot or more high, oblong-lanceolate, acu- minate, lapering, deeply pinnatifid; pinnae oblong-lanceolate, acute, tapering, tomentose tuft at base of each pinna; pinnules obtuse, subcoriaceous, green; veins inconspicuous, veinlets once- forked near midvein; margins entire or obscurely crenulate; fertile fronds about as tall as the sterile, bipinnate, and covered with cinnamon-colored sporangia, arising early in the spring pre- ceding the sterile ones. May and June. Distribution: Newfoundland, westward to Minnesota and New Mexico. Sand- stone. Specimens examined: Eggert, Trelease, Kellogg, Pinkerton, RusselL 3. Osmunda Claytoniana L. Sp. PI. 2: 1066. 1753. Gray, p. 40; Britton & Brown, ed. 1, 1:6, ed. 2, 1:8. Rootstocks creeping, stout, with imbricated leaf-bases; stipes several inches to 2 feet long, woolly when young but never chaffy; fertile fronds taller than the sterile and in the midst of the crown formation, oblong-lanceolate, 1-4 feet long; lowest pinnae about half as long as middle ones, acute and often rounded; pinnae barely acute, never acuminate, short-stalked, lanceolate from a broad base; pinnules close; 2-6 pairs of fertile pinnae near middle of frond, shorter than sterile pinnae and deflexed in a mature speci- men, closely bipinnate, woolly, covered with bivalvular reticulated sporangia. May to July. Distribution: Newfoundland to North Carolina, westward to Minnesota and Missouri. Swamps and moist woods, moist sandstone ledges. Specimens examined: Bush, Eggert, Davis, Palmer. Lycopodiaceae LYCOPODIUM 1. Sporophylls segregated into slender cones; habit of plants fan-like 8. L. complanatum var. flabelliforme 1933] PINKERTON — FERNS OF MISSOURI 73 1. Sporophylls not differing from vegetative leaves; habit of plants rope-like 2 2. All the leaves broadest above the middle; margins jagged 1. L. lucididum 2. Shorter leaves broadest at base; margins entire or slightly denticulate 2. L. lucididum var. porophilum 1. Lycopodium lucidulum Michx. Fl. Bor. Am. 2: 284. 1803. Gray, p. 55; Britton & Brown, ed. 1, 1 : 40, ed. 2, 1 : 44. Stems assurgent from decumbent persistent bases giving rise to a few vertical stems ; leaves dark green and shining, widespread or becoming deflexed, acute, broadest above middle, erose- denticulate, arranged in alternating series of long and short members, the latter often entire and usually bearing the sporangia; gemmiferous. August to October. Distribution: Newfoundland to Delaware, westward to Alaska and Washington. On sandstone only, usually associated with Sphagnum. Specimens examined: Kellogg, Eggert, Pinkerton. 2. Lycopodium lucidulum var. porophilum (Lloyd & Underw.) Clute, Fern Allies, p. 3. 1905. Gray, p. 55; Britton & Brown, ed. 2, 1 : 44. Lycopodium porophilum Lloyd & Underw. in Bull. Torr. Bot. Club 27: 150. 1900. Essentially like L. lucidulum except that the shorter leaves are broadest at the base and the margins are nearly smooth, and are not deflexed. Distribution: in the same places as the species. Sandstone. Specimens examined: Terre Bleue Cr., Ste. Genevieve Co., coll. of Aug. 29, 1898, Trelease, and Pinkerton 81. 3. Lycopodium complanatum var. flabelliforme Fernald in Rhodora 3: 280. 1901. Gray, p. 57; Britton & Brown, ed. 1, 1 : 43, ed. 2, 1 : 48. Rhizomes slender, creeping, with numerous erect stems which branch irregularly, giving rise to a flattened fan-shaped vege- tative structure, about a foot high with 4-ranked imbricated scale-leaves, those of the two lateral rows broad, with spreading tips, of the upper row narrow and incurved, and of the lower row minute deltoid-cuspidate; peduncles long, dichotomously branched at tips and bearing a number of slender cones about an inch long. August and September. 74 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 20 Distribution : Greenland to West Virginia, westward to Alaska and Idaho. Open pine woods on sandy soil. Specimens examined: Pickle Springs, Ste. Genevieve Co., Kellogg 8718. Selaginellaceae SELAGINELLA 1. Plants bearing ill-defined strobili; leaves dimorphic, 4-ranked 1. S. apoda 1. Plants bearing distinct strobili; leaves of one kind, spirally arranged. 2. S. rupestris 1. Selaginella apoda (L.) Fernald in Rhodora 17: 68. 1915. Gray, p. 58; Britton & Brown, ed. 1, 1 : 45, ed. 2, 1 : 49. Selaginella apus (L.) Spring in Mart. Fl. Bras. I 2 : 119. 1840. Stems prostrate and creeping, 1-4 inches long, pale green, delicate; leaves of two kinds, four-ranked and spreading, the smaller pointed and appressed to the stem; no distinct cones; fertile leaves near tip of branches, those containing macrospores conspicuously bulged. July to September. Distribution: Massachusetts to Florida, westward to Michigan and Louisiana. Moist shaded places, among grasses. Specimens examined: Eggert, Hush, Palmer, Mackenzie. 2. Selaginella rupestris (L.) Spring in Mart. FL Bras. I 2 : 118. 1840. Gray, p. 57; Britton & Brown, ed. 1, 1 : 44, ed. 2, 1 : 49. Stems densely tufted, bearing occasional sterile runners; all leaves alike, narrow, appressed, and imbricated, bristle-tipped, gray-green ; strobili four-sided. August to October. Distribution: Quebec to Alabama, westward to Minnesota and Oklahoma. In dry exposed places where there is a little soil, sandstone, or chert. Specimens examined: Eggcrt, h'ussell, Bush, Palmer, PinkerUm, Greene, Broadhead, Vanlngvn, Shepard. ISOETACEAE ISOETES 1. Megaspores reticulate; sporangia unmarked 1. I. Engclmanni 1. Megaspores tuberculate; sporangia marked in some way 2 2. Megaspores less than 480 \i in diameter; sporangia marked with brown spots. 2. I. melanopoda 2. Megaspores more than 4S0 ^ in diameter; sporangia marked with brown lines. 3. I. Butleri 1. Isoetes Engelmanni A. Br. in Flora 29: 178. 1846. Gray, p. 61; Britton & Brown, ed. 1, 1: 48, ed. 2, 1: 53. Conns 2-lobed; leaves 15-60, 13-50 cm. long, light green; sto- 1933] PINKERTON — FERNS OF MISSOURI 75 mata numerous; peripheral strands variable in number or none; sporangia oblong, unspotted, with narrow velum; megaspores white, 400-570 [x in diameter, distinctly marked with honey- comb network of narrow ridges; microspores 21-30 fi, seldom 33 (x, in length, smooth to minutely roughened. Distribution: eastern border to Mississippi valley. Near ponds. Specimens examined: Engelmann. 2. Isoetes melanopoda Gay and Dur. in Bull. Soc. Bot. Fr. 11 : 102. 1864. Gray, p. 61 ; Britton & Brown, ed. 1, 1 : 48, ed. 2, 1 : 54. Corms 2-lobed; leaves 15-60, 15-40 cm. long, slender, erect, firm, bright green, usually black and shining at base, with usually pale membranaceous border, little (2-3 cm.) extended above spo- rangium level; stomata present; peripheral strands 4-6 cardinal, plus as many as 14 accessory groups; ligule subulate, triangular; sporangia oblong, 0.5-3 cm. long, marked by numerous brown spots; velum variable, from very narrow to covering half of sporangium ; megaspores 280-440 p in diameter, marked with low tubercules, frequently confluent into short low wrinkles; micro- spores frequently ashy-gray, 20-30 \i long, fine-spinulose. Distribution : Illinois to Texas. Wet prairies. Specimens examined: Pfeiffer, Bush, Palmer. 3. Isoetes Butleri Engelm. in Bot. Gaz. 3:1. 1878. Gray, p. 61; Britton & Brown, ed. 1, 1 : 48, ed. 2, 1 : 54. Corms 2-lobed; leaves 8-30, 8-15 cm. long, more slender and rigid than /. melanopoda, tapering to apex; stomata numerous; peripheral strands usually 4, sometimes more in number; ligule elongated, cordate at base; sporangium oblong, 6-7 mm. long, marked with brown lines; velum very narrow; megaspores vari- able, commonly 480-650 [x in diameter, marked with numerous tubercules, usually distinct, occasionally confluent; microspores 27-37 [t. long, papillose. Distribution: Tennessee, westward to Kansas and Oklahoma. Limestone barrens. Specimens examined: Eggert, Bush, Palmer. Salviniaceae AZOLLA 1. Azolla caroliniana Willd. Sp. PL 5: 541. 1810. Gray, p. 50; Britton & Brown, ed. 1, 1: 35, ed. 2, 1: 38. [Vol. 20 »T 76 ANNALS OF THE MISSOURI BOTANICAL GARDEN Plants floating on surface of water, often covering large areas, deltoid or triangular-ovate in outline, 6-25 mm. broad, pinnately branched ; lobes ovate, lower lobe reddish, upper greenish with a reddish border; megaspores minutely granulate with three acces- sory corpuscles; masses of microspores armed with rigid septate processes. Washi On surface of still waters. Specimens examined: Eggert, Engelmann, Trelease, Bush, Mackenzie. INDEX TO SPECIES 9 Acrostic/mm arcolatum 69 Adiantum 50 Capillus-Veneris 50 pedatum 50 Aspidium 66 Azolla 75 asplenioides 55 Filix-foemina 55 pycnocarpon 54 thelypteroides 55 acrostichovhs 63 caroliniana 75 var. incisum 64 Botrychium 47 schwdnitzii 64 Asplenium 51 acrostichoidcs 55 angustifolium 54 Athyrinm 55 Bradleyi 51 cryptolepis 51 ebenemn 53 ebenoides 52 dissectum 48 obliquum 47 var. dissectum 48 var. tenuifolium 48 ternatum 47 var. dissectum 48 var. obliquum 47 virginianum 48 var. intermedium 49 Filix-foemina 56 Camptosorus 57 56 rhizophyllus 57 Mid HlUXll parvulum 53 Cheilanthes 57 pinnatifidum 52 platyneuron 53 53 54 53 alabamensis 57 var. incisum dealbata Feci . . . 61 58 lanosa 58 lanuginosa 58 lluta-muraria 52 vestita 58 thelypteroides 55 Cystopteris 58 pycnocarpon resiliens. . . Trichomanes 54 Athyrinm 54 acrostichoidcs 55 bulbif era 59 var. horizontalis 60 fragilis 58 angustifolium 54 Dennstaedtia 60 angustum 56 punctilobula 60 var. elatius 56 Dicksonia 60 var. rubellum 56 9 Names in italics are synonyms. pilosiuscula 60 1933 PINKERTON — FERNS OF MISSOURI 77 punctilobula 60 Dryopteris 66 acrostichoides 63 Equisetum 69 arvense 69 hyemale robustum 70 laevigatum 71 praealtum 70 robustum 70 Filix 59 bulbifera 59 fragilis 59 Hippochaete prealta 70 Isoetes 74 Butleri 75 Engelmanni 74 melanopoda 75 Lastrea 66 Lorinseria areolata 69 Lycopodium 72 complanatum flabelliforme 73 lucidulum 73 var. porophilum 73 porophilum 73 Matteuccia nodulosa 65 Struthiopteris 65 Nephr odium 66 Notholaena 60 dealbata 60 nivea dealbata 61 Nothochlaena pvlchella 61 Onoclea 61 sensibilis 61 Struthiopteris 64 Ophioglossum 46 Engelmanni 47 vulgatum 46 Osmunda 71 cinnamomea 72 Clay toniana 72 regalis spectabilis 71 Struthiopteris 64 Pellaea 61 atropurpurea 62 var. Bushii 62 var. cristata 62 dealbata 61 glabella 62 Phegopteris 66 hexagonoptera 68 Polypodium 62 incanum 63 polypodioides 63 virginianum 63 vulgar e 63 Polystichum 63 acrostichoides 63 var. incisum 64 var. Schweinitzii 64 Polystichum 66 Pteretis 64 nodulosa 64 Pteridium 65 aquilinum 65 latiusculum 65 var. pseudocaudatum 65 Pteris aquilina 65 Selaginella 74 apoda 74 apus 74 rupestris 74 Struthiopteris germanica 65 Thelypteris 66 hexagonoptera 68 marginalia 67 palustris pubescens 66 spinulosa 67 var. intermedia 67 Thelypteris 66 Woodsia 68 obtusa 68 Woodwardia 69 angustifolia 69 areolata 69 [Vol. 20, 1933] 78 ANNALS OF THE MISSOURI BOTANICAL GARDEN EXPIANATION OF PLATE PLATE 5 Fig. 1. Cystopteris bulbifera var. horizontalis Lawson. From E. J. Palmer No. 82896, in the Herbarium of the Missouri Botanical Garden. Fig. 2. Botrychium obliquum var. tenuifolium (Underw.) Gilbert. From Dr. William Trelease, coll. of October 28, 1897, in the Herbarium of the Missouri Bo- tanical Garden. Ann. Mo. Bot. Gaud., Vol. 20, l!i:i3 Plate 5 -o X O ! ► c *1 C w BLASTOMYCOSIS: REPORT OF A CASE, WITH A STUDY OF AN ETIOLOGIC FACTOR AND A CLASSIFICATION OF THE ORGANISM MORRIS MOORE Rufus J, Lackland Research Fellow in the Henry Shaw School of Botany of Washington University Introduction It is the purpose of this paper to describe briefly the disease known clinically as blastomycosis, and to try to clarify the rec- ognition of the organism involved. The literature of the field is at present too extensive for an entire review, and since numer- ous workers have already given excellent discussions on the clinical aspects of the infection, as to its gross pathology, micro- scopic histo-pathology or cellular reactions, and the biological or rather immunological phenomena, a review would be un- necessary here. However numerous such papers may be, there is still much work to be done on the subject. The author has attempted to clear up, at least in his own mind, several undecided points in the disease: first, the establishment of the etiological agent of blastomycosis; second, the determina- tion of the exact classification of the organism. In the past, and even at the present, medical men have grouped under one general heading all organisms which were responsible for the same clinical condition. Good as this system may be for general diagnosis, much difficulty is encountered because of the fact that physicians are inclined to devote very little time to a study of the organism, thus rendering any therapeutic measures, if avail- able, indefinite, inasmuch as several of the fungi present varying degrees of pathogenicity and require different therapeutic meas- ures. Thus we find that numerous species of the genera Saccharo- myces, Monilia, Cryptococcus, Endomyces, Sporotrichum, and others have, at one time or another, been considered etiological agents of blastomycosis. A review of the history will illustrate these facts. History Years before the first case was definitely described as blastomy- had performed a certain amount of Ann. Mo. Gard., Vol. 20, 1933 (79) 80 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 20 fungi involved in cases of infection and had established these organisms as etiological factors, particularly the yeast and yeast- like groups. Chronologically, the list is quite long, but it is worthy of note. According to Hufschmitt, Sartory, Sartory, and Meyer ('31), we find that in 1845 Remak, and in 1853 Robin, in his 'Histoire Naturelle des Veg6taux Parasites de l'Homme et des Animaux/ discovered the normal existence of the yeast Cryptococcus guttu- latus in the rabbit intestine. A few years previously, Hannover (cited in Buschke and Joseph, '28) had found yeast in the urine of diabetic patients. Investigations then tended to turn to the parasitism of these organisms in animals, with the result that Bernard during the course of his work on fermentations attempted the first animal experiment by injecting beer yeast into these subjects. Following this work, Popoff, Grohe, Roussy, and several others showed the pathogenic actions of the yeasts on mammals, and Rivolta in 1873 in his 'Parasiti Vegetali' dem- onstrated lor the first time a yeast infection in a horse. In the meantime, Metchnikoff and Weismann showed the parasitism of the Saccharomycetaceae in the lower animals. In 1892 Wernicke showed the first mycosis in man and named it "maladie protozoique de la peau." The following year Troisier and Achalme ('93) definitely established the relation between yeasts and man. In the meantime, several workers attempted to show the destructive ability of these organisms on the animal tissues. Popoff in his work had used dogs as his subjects, but impure cultures. Rauni ('91) inoculated animals with large amounts of yeasts, and a rise in their temperature, shortness of breath, and death resulted. Neumayer ('91) fed animals with cultures and also inoculated them subepidermally. His feeding developed a gastro-enteritis which he believed due to fermentation, since the skin inoculations were of no value. The yeasts of these workers were probably of the non-pathogenic types, for L. Rabinowitsch ('96), a few years later, showed fifty various yeast-like organisms with seven pathogenic for animals. Nesczadimenko ('99) made peritoneal injections of yeasts in a physiological saline solution into rats, mice, guinea-pigs, and dogs, with death ensuing from eight to twelve days. He concluded, however, that these organisms were not so deadly, although causing this mortality. 1933] MOORE — BLASTOMYCOSIS 81 The first actual case of blastomycosis, so-called, was reported by Gilchrist at the June, 1894, session of the American Dermato- logical Association. His paper resulted from the finding of peculiar yeast-like bodies in the diseased tissue of a patient. The doctor attending the patient had given the diagnosis as a typical case of chronic scrofuloderma. Several months after Gilchrist's report, Busse ('94) brought to light the extraordinary case to which he later gave the name " Saccharomycosis hominis." The patient was a woman thirty-one years of age who had suffered from a localized subperiosteal inflammation of the left tibia. An examination of the abscess, which opened spontaneously, re- vealed "numerous doubly contoured, very refractive, roundish and ovoid bodies," and these were found to be situated both intracellularly and extracellularly in the pus and abscess wall. These organisms when isolated in pure culture and then inocu- lated experimentally in animals proved to be what were later known to be blastomycetes. The patient later developed super- ficial ulcers on the face, subperiosteal swellings on the right ulna and the left sixth rib near the axillary line, with death ensuing. Busse cultured the yeast from the ulnar swellings and from the bottom of the ulcers. Approximately two years later, the first case reported by Gilchrist was published in detail in the Johns Hopkins Hospital Reports of 1896. In the meantime, however, several others had noticed similar cases among guinea-pigs, horses, mice, and other lower animals (Sanfelice, '95, '96, '96a, Roncali, '95, Corselli and Frisco, '95, Tokishige, '96, and others). In 1896, Curtis isolated a fungus similar to that described by the former writers from a myxomatous tumor of the leg. In the same year, Gilchrist, in conjunction with Stokes, published a short paper on a second case of blastomycosis, and this was published in detail two years later (Gilchrist and Stokes, '98). Simoni ('97), working on the diseased tonsils of patients, found budding Maffuci and Sirleo '98) examined numerous tumors and found budding cells in a great number of tissues. Many other reports followed, as that of Hyde, Hektoen, and Bevan ('99) with a supplement by Hek- toen C'99^) later in the vear. Hessler C99) with a case report, and 82 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 20 several during the same year and 1900. In the following year appeared the elaborate work of Ricketts ('01), with a study of the organism from a case of systemic blastomycosis by Otis and Evans ('03). Eisendrath and Ormsby ('05) described a systemic infection, and Irons and Graham ('06) reported a severe gener- alized systemic disorder. Hektoen ('07) gave a comprehensive review of the literature, and from that time on the medical journals have published too great a number of cases of infection due to yeast-like organisms, under the heading of blastomycosis, to render a complete survey of literature a matter for a paper of this length. Etiology and Clinical Manifestations The disease known clinically as blastomycosis is very likely due to a plurality of organisms and not species of the same genus as indicated by previous writers. This is clearly evident as seen by the great number of papers published and the cases reported, involving such fungi as Saccharomyces, Oidium, Monilia, En- domyces, Cryptococcus, C occidiaides , and even such a form as Sporotrichum. The clinicians have referred to the category of blastomycosis any clinico-pathological condition which may be due to yeast-like or budding fungi. It must be understood, therefore, that the term as here used refers only to the clinical aspect of the condition. In America, clinicians and medical men, more especially medical mycologists, are inclined to class as the cause of blas- tomycosis only that organism which was described originally by Gilchrist and Stokes in 1894 and in this view the author is greatly in accord. On the other hand, European workers consider only that organism which was reported by Busse ('94) and so elaborated on by Buschke ('98). However, by reason of priority, the organism of the former workers should hold the position so designated and be established as such. Further remarks on the Gilchrist organ- ism will be found in the discussion. Blastomycosis presents numerous clinical manifestations and in this respect it is protean, being found in practically every organ of the human body either in biopsy or autopsy material. No immunity towards the invading organism is established by any 1933] MOORE — BLASTOMYCOSIS 83 of the anatomical structures. Clinically, the condition presents lesions which are alike both for the cutaneous type of the disease, that is that group of infections which may be found occurring superficially, or for the systemic type of the disorder, occurring in the lungs, bones, meninges, liver, or other viscera. This division is based on the work of Jacobson and his associates ('32), who further separate the cutaneous type into that of primary in character, as occurring in the epidermic layers or the cutis, as shown by Hagiwara ('22) and Hashimoto ('22), whose organisms, although not of the Gilchrist type yet coming under the general heading of blastomycosis as known clinically, also the work of Grschebin ('27) ; and secondary, due to an infection of the deeper tissues, internal viscera, or bony structures, as shown by Irons and Graham ('06), Ryerson ('09) for bones, and many others. The primary form or the cutis infection Jacobson further designates as presenting one of three varied appearances : papulo- ulcerative; verrucous or papillomatous; gummatous. The papulo-ulcerative type Jacobson designates as being initial lesions which are papulo-pustular in character and of epidermic origin (shown by Hessler, '99, Hektoen, '99, Ricketts, '01, Engelhardt, '24, and Fabry, '27, '28). These lesions rupture in the course of time and empty out the purulent exudate on the surface of the skin, with the probable ultimate formation of crusts. The process may be proliferative and involve a great area of the immediate vicinity. The lesions usually show a violaceous border with the involution of the peripheral surfaces and perhaps consequent scarring and atrophy. The verrucous or papillomatous type (Froilano De Mello and Rodrigues, '29) is characterized as being nodular or papular in character and present on a normal or deep-red, infiltrated skin. Several of the lesions may coalesce to form papillomatous patches which resemble verrucous tuberculosis. These lesions may break up into healing areas which upon drying present irregular scars. The characteristic color as noted above is found here too, as well as the sloping periphery. The gummatous type develops from the subcutaneous layers of the tissue of the deeper portion of the cutis in the form of small, [Vol. 20 84 ANNALS OF THE MISSOURI BOTANICAL GARDEN slightly elevated, somewhat tender, reddish, deep-seated, soft nodules situated on the characteristic violaceous-red surface of the skin. There is a diffusion of the color with subsequent establishment of new nodules in the vicinity. The nodules en- large, become soft and gummatous, and then break down to form masses of ulcerative, proliferative materials bordered as in the other two types, and contain numerous abscesses. The secondary cutaneous form consists chiefly of variously formed ulcers which give off a purulent or sanguino-purulent discharge from a soft, granulating floor. Some may develop crusts with raised edges, while some may assume hyperplastic functions with papillomatous characteristics, and usually there may be a metastatic action on the part of the ulcers represented by the formation of new lesions which are surrounded by a dark red or purplish zone. Healing may be spontaneous with indurated scars, as noted in some cases, or infection may persist but may finally succumb to treatment with iodides as was noted in the case reported here. A study of the ulcers formed in blastomycosis shows them to originate in abscesses which from a clinical point of view can be divided into the superficial and the deep types. Secondary, cutaneous, superficial ulcers arise usually in the subcutaneous tissues aa nodules of varying size as shown by Stober ('14), Engelhard t ('24), Ferguson ('28), and Montgomery and Ormsby ('08). These ulcers usually enlarge, rupture, and spread the material over the surface of the skin, setting up new foci, or in some cases they have been found to dry up and disappear. The deep type of secondary cutaneous blastomycosis (Grsche- bin, '27, '28), characterized by smaller number and deep- seatedness, is by far the most serious of the two, involving de- structive processes of the bone, muscle, and deep tissues and organs. It rarely shows any inflammatory reaction, but can be distinguished by the purulent character of the abscesses as con- trasted with the mucoid or mucopurulent nature of the super- ficial abscesses. The above types represent the typical forms occurring in a clinic. However, Weidman and Douglas ('21) reported the occurrence of a sarcoma-like tumor on the lee of a patient, which 1933] MOORE — BLASTOMYCOSIS 85 looked like lupus vulgaris and yielded blastomycetoid bodies on histological sectioning. Then, about six years later, Cleland ('27) reported a case with the formation of a myxomatous-looking tumor mass which also showed tvoical cells on sectioning. typical cells on sectioning. These, however, are rare and until more cases are reported cannot be placed in the definite clinical types. Under the heading of cutaneous blastomycosis, Castellani has also established principal types of blastomycosis of the cutis from clinical Doint of type) Etiology: Cryptococcus dermatitidis Gilchrist and Stokes, 1896 (Synonym: Cryp- :occus gilchristi Vuillemin). Castellani here creates the genus Blastomycoides . 2. Blastomycosis ulcerativa profunda sen mutilans (Synonym: Blastomycosis, Wernicke Coccidioides 3. Blastomycosis purulenta profunda (Synonym: Blastomycosis, Busse type; Blastomycosis subcutanea purulenta). Etiology: Cryptococcus hominis Vuillemin, 1901 (probably covers many species). 4. Blastomycosis glutealis (Synonym: Blastomycosis, Kartulis type). Etiology: mycological investigations not yet completed. The fungi seem to belong to the genera Monilia and Cryptococcus. 5. Furunculosis bl ostomy celica cryptococcica (Folliculitis decalvans cryptococcica, pro parte, Castellani type). (Synonyms: Furunculosis cryptococcica; Pseudo- furunculosis blastomycetica; Furunculosis mycetica; Folliculitis decalvans sacchar- omycetica; Folliculitis decalvans moniliaca). Etiology: yeast-like fungi either of genus Cryptococcus or Monilia (No asci or ascospores according to Castellani). In addition to the above types, Castellani adds the following, although they are in no way connected with the blastomycosis organism: Blastomycosis epidermica; Intertrigo blastomycetica; Dermatitis blastomycetica interdigitalis. For clinical purposes in diagnosing skin infections such a classification is good, but for correct determination of the etio- logical agent, it is essential that each type of involvement be named with the infective agent designated as such. For example, if the organism be a saccharomycete, the disease should be called saccharomycosis; if a monilia, then moniliomycosis; if an endomy- cete, then endomycosis. Thus, when the organism is correctly diagnosed the amount of time necessary to determine the right sort of curative measure for that type of infection maybe taken, and the amount of time necessary for healing reduced. [Vol. 20 86 ANNALS OF THE MISSOURI BOTANICAL GARDEN Systemic Blastomycosis As mentioned previously, the disease is protean in its clinical manifestations, with the result that practically every organ in the body has been shown to be infected either in the living, by various measures, such as X-ray, or in autopsy material. No vital organ is immune, and this in itself is sufficient to cause a careful physician to give more attention to therapeutic measures. This universal infectivity of the agent was especially shown by such writers as Otis and Evans ('03), Eisendrath and Ormsby '05), Le Count and Myers ('05), Irons and Graham ('06), Montgomery and Ormsby ('08), Wade and Bell ('16), Garr ('25), Panja ('25), Toepel ('29), and Maner and Hammack ('30). There is, however, a difference in frequency wit h which the vari- ous organs show their susceptibility. The port ion of the anatomy that shows the greatest amount of infection is the skin, either primary or secondary, having about 95 per cent of all cases recorded. This phase of blastomycosis has received the greatest amount of attention principally because it is so prevalent, but also because it is usually a manifestation of a metastasis from the deeper organs, and this helps bring forth the diagnosis of blastomycosis. The pulmonary system, including the lungs and bronchi, constitutes the second most frequent and the most common sys- temic form, being present to theextentof approximately 95 percent in systemic infection in available autopsy records, and about 35 per cent in primary cases. This was shown by such menas Stober ('14), Wade and Bell ('16), Wade ('18), Dennis ('18), Miller ('27), Medlar ('27), and Mazza and Nifio ('28). The disease probably is primary in the bronchi and from there spreads to the lungs. If secondary in the lungs, as in systemic disorders, the process may be slow and chronic; if primary, however, the spread may be rapid and fatal. The kidneys are next in frequency of infection. The genito- urinary involvement is usually secondary by way of metastatic foci through the blood. The disease in these organs manifests itself in the form of nephritis, showing casts and albumen in the urine. The culturability of the organism from samples, however, 19331 MOORE — BLASTOMYCOSIS 87 cannot be demonstrated unless the kidneys are accompanied by infected bladder or prostates. The spleen follows next in order, but this organ is usually easily susceptible so that a great amount of infection is to be expected in any systemic disorder of this sort. The complication of the bones and joints seems next in the amount of infection. This form of the syndrome is very common in systemic disorders, and may even be a primary infection if the diagnosis in the patient here described was correct seven years previous to his entry at the Barnard Skin and Cancer Hospital. This type of the disease may manifest itself in the form of arthritis, osteitis, osteomyelitis, or periostitis, according to Ryerson ('08-'09) and Stober ('14). The process usually results in a suppuration, formation of sequestra, and abscesses which break down intervening cell walls and coalesce, causing great damage. The liver appears to be a rather frequent subject to the in- fection, coming next in the order of frequency. This is to be expected in systemic disorders where the blood plays an impor- tant part. Metastases through the blood vessels are fairly com- mon, and yeast cells are easily transported to the main organs in this manner. The pleura too are susceptible to a great extent, and here the proximity to the lungs is a great factor in their infection. The lymph glands follow shown by Wanamaker ('28), especially for the cervical lymph glands. Cerebrospinal involvement, including the brain, meninges, spinal cord, and skull bones, occurs fairly often as a secondary metastatic process in generalized systemic blastomycosis, ac- cording to studies made (J. T. Moore, '20, Freeman and Weid- man, '23, Greenfield, '24, Wilhelmj, '25, and Gaspar, '29), being found When the disease secondary to a general systemic infection, there may be osteo- myelitis of the skull bones with destructive processes. The diagnosis rests not upon any clinical entities which may be present, because the inflammatory reaction simulates many other condi- tions, but upon the actual laboratory finding of the organism either in the spinal fluid or in sections of the diseased brain tissue. [Vol. 20 88 ANNALS OF THE MISSOURI BOTANICAL GARDEN Wilhelmj ('25) states that in those cases where there is no patho- logical symptomatology or clinical manifestation on any other part of the body, and when the meninges are infected during the primary stages of the invasion, death may occur without the initial appearance of general metastatic foci, and such a condition he calls primary cerebrospinal blastomycosis. The spine may be involved in the process in a suppurating condition (Parker, '23), but this condition is relatively rare. Jacobson lists the vertebrae as being next to the brain in susceptibility to attack. This condition has been noted on several occasions. Roentgenographic studies usually reveal an infection of bodies of the vertebrae, and lamina and posterior ligaments may show an involvement which simulates greatly tuberculous Pott's disease. Prostatic infection in blastomycosis is often noticed (Parmenter and Simpson, '19). Usually it is associated with a genito-urinary complication and involves the urinary bladder (Rhamy, '26). In these cases acute urinary urgency and pyuria are well-defined symptoms. The heart lesions in blastomycosis are shown first in the pericardium and then in the myocardium in the form of an in- flammatory reaction (Hurley, '!()). Pancreatic involvement follows in frequency. Infection in the peritoneum is the next most common. Jacob- son reports finding the disease in the abdominal viscera in the following decreasing order of frequency: kidneys, spleen, liver, lymph glands, pancreas, peritoneum, adrenals, and gastro- intestinal tract. These organs, as pointed out previously, become involved usually through metastasis by way of the blood stream or by direct transmission from tissue to tissue. In this manner, testicular blastomycosis is usually developed. Planchard, Swartz and Binot ('03), as early as 1903, noted an intraperitoneal in- volvement. The eyes may also be involved. The infection here is very painful and often dangerous, leading to blindness with perhaps a complication of the nervous system and the brain. McKee ('26) and Ferguson ('28) noted cases of the eye which were secondary infections due to a metastasis from the pulmonary apparatus. 1933] MOORE — BLASTOMYCOSIS 89 Laryngeal and tracheal blastomycosis are rare infections. Jacobson records four in America and one in Europe. All the patients were adult males working either with the soil or its products, one being a clerk in a general store (Dennis/ 18, Downing, '18, C. Jackson, '26, and New, '28). The larynx showed a ' ' chronic inflammatory mucosa with a grayish, minutely nodular surface in some portion of the lesion, with a few minute, isolated, yellowish nodules." There was often a reddish, raw portion of the larynx due to the ease with which the superficial layers came off with coughing. The process resembled very closely tuberculosis. Involvement of the tongue is perhaps a rare occurrence, but cases are not reported in great numbers purely because sputa smears usually show a variety of yeasts and thus no definite etiological significance is attached to those obtained. The first case was reported by Copelli ('13), a second one by New ('17) from the Mayo Clinic. Since that time, however, Mazza and Canevari ('29) reported a case from Argentina, and Nino ('29) reported an infection of the lower lip with the involvement of the tongue. Such an infection usually hinders respiration, inasmuch as a tumor-like growth, as evidenced by Copelli and New, de- veloped which enlarged in the back portion of the tongue and practically filled the entire larynx. numerous Symptomatology Blastomycosis when of the primary cutaneous type presents no clinical symptomatology of discomfort or pain except for that expressed because of the lesions. When of the systemic type, however, the condition is very different. There are clinical factors to make a picture which might easily be confused with a number of diseases. According to numerous investigators, the onset of the disease varies with the person and amount of infection. It may be intense and acute, leading to death in a short time, or insidious and mild, with a prolonged chronic condition, death finally occurring as a result of a secondary complication. There is a typical set of symptoms once the disease is well established. This consists of typical malaise, recurrent chills, loss of weight, [Vol. 20 90 ANNALS OF THE MISSOURI BOTANICAL GARDEN as evidenced in the present case, loss of strength leading to general emaciation, night sweats, although morning sweats may be present too, irregular fever, pain in affected parts, and a rapid pulse. The disease, as noted before, may be primary in the skin with subsequent spread systemically or it may be systemic with the formation in the later stages of nodular growths on the skin. Unless the patient is well taken care of, systemic infection re- sults, leading ultimately to death. Differential Diagnosis The final diagnosis of blastomycosis rests on the finding of the organism either culturally in a lesion of the patient or, if that be unavailable as in systemic disorders, the identification of the fungus in biopsy or autopsy material. As pointed out previously, the disease is protean in character, with the result that a careful examination must be made to diagnose it blastomycosis, comparing it with the several well- known clinico-pathological entities which it may simulate. The most noteworthy of these complicating diseases are as follows: (a) The dermic lesions described previously, developing necrotic and papillomatous growths or ulcers, resemble very closely epitheliomas, differing only in the rapidity of evolution and the absence of deep induration, verrucas, tuberculosis in its various forms, and syphilis. Its resemblance to sporotrichosis (Lewis, '17) has often been noted, but it differs in being less sluggish. It differs from syphilis only by the softness of the lesions, the red- dish-blue ring around the lesion, and by a negative Wassermann, with, of course, the presence of the organism in the blastomycosis infection; (b) The systemic infections of blastomycosis must be distinguished through laboratory methods from a great many complications, particularly coccidioidal granuloma described in a previous paper by the author (M. Moore, '32). The organism of coccidioidal granuloma, Coccidioides immitis, reproduces by en- dosporulation, and the blastomycosis organism, through budding. Furthermore, the lesions in the so-called "California disease" are more rapid in evolution than in blastomycosis; (c) Infection of the glands which is quite rare in blastomycosis often suggests 1933] MOORE — BLASTOMYCOSIS 91 lymphatic leukemia, Hodgkin's disease, and possibly lympho- sarcoma; (d) Gastro-intestinal lesions may often resemble typhoid and in some cases the isolation of the organism is necessary to rule out this disease; (e) Osseous infection quite often resembles tuberculosis, particularly as reported by Ryerson ('08-09); Pulmonary blastomycosis very often presents the same clinical, histological, and pathological pictures as tuberculosis, as noted by Medlar ('27) and Miller ('27) ; (g) Infections of the brain often confuse the pathologist or clinician with its similarity to torula (Freeman and Weidman, '23), epidemic meningitis, and even tumors of the spinal cord; (h) Sutejew, Utenkow, and Zeitlin ('29) find that the use of bromides and iodides evidently causes an allergy which in its reactions presents lesions similar to those caused by the infective agent of blastomycosis and is often confused with it, the latter differing in their more rapid evo- lution. It would seem, therefore, that the recognition of blastomycosis is not a very easy matter. The really important fact concerned with this work is to find the organism, which usually requires laboratory technique, and to verify its pathogenicity by animal inoculation in order to comply with Koch's postulates. Predisposing factors. — The infective agent shows no particular preference as to sex, although more cases have been males than females, of the industrial classes, chiefly the workers of the soil and its products, a fact well exemplified in the present case. There is no discrimination as to race or color, all peoples being affected in like fashion. Stober ('14) correlates the type and amount of infection with the habitat and environment of the r patient. As far as is known, all ages are susceptible, a similar condition existing for many other fungi which tend to become systemic, such as coccidioidal granuloma. Treatment. — The successful therapeutic measures in blastomy- cosis are few and limited. The best treatment or cure for the disease rests of course on the skill of the physician in detecting it at an early stage before it has seized a definite foothold, when it can be kept from becoming systemic. If the disease is definitely located in a particular section, surgery may be used to eliminate it, as suggested by several authors (Wade and Bell, '16, R. H. 92 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 20 Jackson, '2G). Cautery has also been used with beneficial results. Hedge ('28) has employed carbon dioxide snow in freezing cutaneous lesions with measurable success. X-ray treatment has also been used frequently, being combined with the administration of iodides. On the whole, primary cutaneous lesions yield fairly readily to iodides and even the application of crystal violet and gentian violet, although their use if at all successful is empirical because dye therapy does not rest on any scientific basis (Spring, '20). In systemic infections, however, one has to contend with complications. The extent of the infection cannot be determined easily. The only cases to recover from an infection of this sort are those to which special care and attention coupled with a change to a clean, pure atmosphere had been made, with the administration of large amounts of saturated iodide solution, either potassium or sodium, although the former has been used more extensively. The dyes mentioned above have proven worthless to the systemic type of disorder. Several workers have advocated copper sulphate, but many others have found this to be useless. Roentgenotherapy has, as yet, no really therapeutic importance. Stober has applied immuno-therapy in the form of a vaccine of suspended blastomycotic cells, heated to 110° C, but no definite results can be shown. It would seem, therefore, that therapeusis is greatly in need of investigation. Immunological reactions. — Immunology in blastomycosis has not reached any definite point as yet. Agglutinins have been reported by some, and negative results by others. Precipitins have also had the same reaction, as well as complement fixation. On the whole, results are indefinite and a good deal more in- formation is needed. The main difficulty seems to be that the toxins of the blastomycosis fungus are difficult to demonstrate. So far, immunology is an open book with only attempts proving nothing written on its pages, and it is to be hoped that more work can be done along this line for the benefit of those who may be inflicted with this syndrome-complex. Mortality. — The number of deaths resulting from blastomycosis is a factor worthy of note, since systemic disorders due to the 1933] MOORE — BLASTOMYCOSIS 93 typical blastomycotic organism usually prove fatal. This is so because systemic infections are rarely diagnosed as such until there have been cutaneous outbreaks, with the result that thera- peutic measures are given too late for beneficial results. It is difficult to quote figures because there are many cases occurring which have rapid recovery and the physician does not report them. Moreover, true blastomycosis is difficult to diagnose un- less smears and cultures are made from the abscesses. In many instances, several attempts are necessary before any fungous growth is obtained, and unless the investigator is well trained in mycological technique and in the recognition of such forms typical of the Gilchrist organism the application of Koch's postulates would be essential, particularly with mice and guinea-pigs. Report of Case Clinical History. — M. H. L. (Hospital No. 50095), a white, widowed male, farmer by occupation, 43 years of age entered the Barnard Free Skin and Cancer Hospital, at Saint Louis, Missouri, April 5, 1932, with an ulcerating, proliferating infection of the left hand and forearm, which the patient claims to have had for 5 years. Family History. — Father died as a result of high blood pressure, hypertension, at age of 56 years. Mother dead due to throat trouble, at age of 30 years. Three brothers and one sister dead due to an infancy cause. Patient has 7 children, all living and well. Wife dead due to double pneumonia at age of 38 years. No history of diabetes, nephritis, syphilis, or cancer in family. Past History. — Past health has been good; usual childhood diseases. He had pneumonia at 15 years of age, bronchial pneumonia at 18 years of age, and influenza in 1918. He has had no operations other than those to be mentioned in the present illness. Right leg injured 7 years ago. Personal History. — Patient denied any venereal disease. Present Illness. — Patient stated that 7 years previously, a limb had fallen on his right lower leg, causing a knot to form. The physician who examined his leg thought he had a periostitis and opened the lesion on the leg. About 2 years later, or 5 years previous to entering the hospital, a mossy, verrucous-like lesion similar to that on the leg appeared on the back of the left hand. Both the lesions on the right lower leg and on the hand continued to spread. The patient went to the Vanderbilt Univer- sity Hospital, at Nashville, Tennessee, about a year later, at which time the eruption had spread over most of the anterior and lateral sides of the right lower leg with two lesions of the right thigh above the right knee and with involvement with most of the back of the left hand and part of the forearm. He remained in the Vanderbilt Hospital for about 3 months, during which time the lesions on the right leg and arm were curetted and treated with X-ray. At the time of his discharge, the lesions on his right leg had healed, as had most of the hand except that portion near the thumb and the wrist. Another currettement was performed, but the lesion per- sisted. He received two X-ray treatments each on the leg, arm, and hand. Six years previous to entering the Barnard, he had lesions on the right elbow region; 94 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 20 these lesions healed. About 4 to 6 years before his entry he had lost some weight, but at the present time complains of no recent loss of weight. He gives no history of hemaptysis, night sweats, fever, or frequent colds. About 10 days previous to entry, a reddish lump developed on the flexor surface of the left lower arm near the elbow. This lump is not very tender or painful. He has been applying potash on the ulcerated area of the left hand. Physical Examination. — Patient is a thin, cooperative, moderately active, white male 43 years of age, who has had the condition herein to be described as blastomy- cosis of the left arm and hand for approximately 6 years and who had the same con- dition on the leg for 3 years until it was curetted and X-rayed 4 years previously. The right leg and left arm show scarring which will be described later. Head. — Normal size and shape. Eyes. — Eyes react to light and accommodation. Reactions normal. Ears. — No discharge, apparently normal. Nose. — Septum intact, no ulcers. Throat. — Slight redness. Mouth. — Several teeth missing. Neck. — No stiffness or rigidity; tonsillar and cervical glands palpable. Thorax. — Thorax hairy, thin, and symmetrical; expansion fair. Lungs. — Breath sounds a little harsh and rough over both lungs, principally the right and right apex. Voice sounds normal, but louder on the right than on the left. No persistent or moist r&les or r&les heard after coughing. Expansion and resonance normal. Cardiac. — Cardiac sounds a little slow and distant, but of a normal quality. No enlargement or pathological murmur. Blood pressure, 106 systolic and 66 diastolic. Abdomen. — No masses or tenderness. Inguinal glands a little enlarged. Genitalia. — Normal male; no discharge or penile sores. Reflexes. — Superficial — present and active. Babinski negative. Deep — present and active. Extremities. Left arm and hand — The lower third of the left forearm, back of the hand and extending 2 or 3 cm. on to the palm, is involved in an atrophic process sharply defined superiorly and inferiorly with some scaling, no telangiectasis. On the external surface of the forearm extending on to adjoining parts of the hand and thumb is an ulcerative process which has been covered with a black crust. Ul- cerated area of the left thumb and the flexor surface of the left forearm consists of warty-like and cone-like, multiple abscesses with some elevation of the borders. There is a deep scar on the right elbow region. The anterior, medial, and lateral aspects of the right lower leg are covered with a thin, smooth scar about 12 to 14 inches long and covering two-thirds of the right lower leg. There are two scars just above the right knee, about 4 inches, and 8X4 inches in diameter, respectively. On the flexor surface of the left forearm, just below the elbow, is an abscess which is red in color, oval in shape, semifluctuant, practically non-tender and not hot. Extending up from and about the abscess is a chin of firm lymph nodes. The lymph nodes above the left elbow and epitrochlear region are a little enlarged. Laboratory Findings. — Urine negative, being pale straw in color and clear. Blood tests showed 4,600,000 red blood cells and 10,200 white blond cells, with 84 per cent hemoglobin. The differential blood count showed 26 lymphocytes, 8 large mononu- clears and transitionals, 156 polynuclear neutrophils, 2 polymorphonuclear eosino- phils, and 2 basophiles. Wassermann negative. Smears from the left hand and 1933] MOORE — BLASTOMYCOSIS 95 arm showed budding forms of yeast cells on April 7, 1932. Blood, urine, and pus from left hand and lower arm and pus withdrawn from abscess of left arm were inoculated on glucose-glycerine agar on April 8. Spinal puncture on April 12 showed a clear fluid with a normal pressure. Glucose-glycerine agar and blood agar were inoculated with spinal fluid, w r ith no growth occurring. April 9, 1932. — The patient was started on potassium iodide with dosage up till signs of intoxication, and an ointment saturated with gentian violet was applied locally. This treatment was followed with 10 per cent sodium iodide intravenously for several days when patient showed an improvement. April 16, 1932. — X-ray of left forearm and wrist, right femur, and right leg showed no abnormal bone changes. The hilar and bronchial structures of both lungs showed considerable thickening, inflammatory in character, but the parenchymatous portion of both lungs appeared free from any active pathology. July 7, 1932. — Patient showed a very marked improvement, with still some evidence of trouble around the left thumb. Clinical Diagnosis. — Blastomycosis of left hand and arm. The agar inoculated from the left arm developed a culture which went through the three stages typical of the organism of blastomycosis: the yeast-like growth; the prickly type of growth; and the cottony type of growth. This culture was used for the studies carried out in this paper. Animal Inoculation A suspension of a 10-day-old culture of the organism in sterile saline was inoculated in a mouse, intratesticularly. An orchitis developed and the mouse showed typical malaise, emaciation, loss of appetite, loss of weight, rise in temperature, with death ensuing in three weeks. The internal viscera showed numerous, pin-point lesions at autopsy, which when squeezed exuded a muco-purulent material from which the yeast cells were isolated. This was in accordance with Koch's postulates. Technique In order to ascertain the morphological characteristics of the fungus, the organism was suspended in hanging-drop cultures, allowed to grow, and observations made from time to time. For finer detail and structure, several transfers were made to slides on which had been placed a drop of a mixture of glycerine (Merck C. P.) and a 1 per cent solution of crystal violet. The fungus was allowed to remain for a period of one-half to one hour to allow for a clearing of the material and sufficient staining to render satisfactory results. This method proved adequate for the work here described. Another method was used also, whereby material was fixed on a slide smeared with albumin and then stained with [Vol. 20 96 ANNALS OF THE MISSOURI BOTANICAL GARDEN methylene blue and eosine. The first method, however, was sufficient. DESCRIPTION A study of the organism in lesions reveals a yeast-like growth of spherical or ovoid, budding and branching cells with no evidence of any filamentous hyphae. These cells, measuring approxi- mately 7-12 [i in diameter and sometimes as much as 20 (x in length, may occur singly, in groups of two's, three's or four's, as individual colonies. On closer examination microscopically, these yeast-like cells (pi. 6, figs. 1-9) show a heavv. reticulated mner granular, and in many cases, vacuolated, protoplasm, and a very definite nucleus with emanating streams of cytoplasm. In many cells the nucleus is barely distinguishable, being a mass of cyto- plasmic structures, but further study after several subcultures shows up this part of the cell very frequently. Many of the yeasts of this group may show a double-contoured, highly refractile membrane, and this feature is of little diagnostic use unless dem- onstrated in tissue sections, as by Rewbridge, Dodge, and Ayers '29) for Endomyces capsulatus, and by Moore (McBride and Thompson, '33) for E. capsulatus var. isabellinus . Thus Wade ('10) ascribes to the fungus structure in tissues: (1) an delicate capsula vera; and (2) an outer applied capsula sclerotica. In any case, the capsule is lost on repeated cultivation on arti- ficial media. With agar as a substrate, the yeast-like cells elongate (pi. 6, figs. 10-12, 14-17) and on acid media become thin-walled and long with a diameter of 2-2J^ (x. These hyphae become inter- twined and are composed chiefly of isodiametric cells. On neutral or slightly alkaline media, with protein as the chief source of nitrogen, the hyphae are thicker and shorter, with a diameter of 3-4 [x. This condition is especially true of media with an excess of carbohydrate as found in glycerine agar. Budding cells are numerous on slightly acid media, being about 5 ix in diameter (pi. 6, figs. 26 and 28). The hyphae, at first clear, become granular, and at various points along the sides develop numerous small, knob-like pro- jections of the limiting membrane, which enlarge, round out or become pyriform and sessile, measure approximately 5 u. in 1933] MOORE — BLASTOMYCOSIS 97 diameter, and occur usually near a septum. These are known as conidia and may remain attached, at times to small stems or sterigmata, or may become free and develop in the media by budding (pi. 6, figs. 24, 26, 28-29). Racquet mycelium (pi. 6, figs. 23-24), a phenomenon char- acteristic of the fungi of this group and especially of the Tri- chophytons and various other Ascomycetes, is common here, having the swollen portion 5-6 [/. in diameter and the narrow section 3-33/2 v. in diameter. Chlamydospores may be found arising in the hyphae or terminal as hypnospores, varying in size and shape from round cells 7-83^ [l in diameter, to elongated, widened cells 53^-7^ ;x wide and 12-15 jx long (pi. 6, figs. 22, 25, 27-29, 31, 32, and 37), or they may arise as sessile cells from the hyphae (pi. 6, fig. 36). Round, thick cells with a coarse, granular cytoplasm are particularly evident on cornmeal agar (pi. 6, figs. 26 and 28). When first examined in a hanging-drop preparation, one may see oil droplets on the hyphae which are strongly suggestive of endospores. These disappear, however, when the organism is stained. In some cases, as in pi. 6, fig. 25, a hollow sphere or vacuole surrounded by a hyaline, gelatinous substance may be present in the filament. The development in tissue, as has been noted, is chiefly by budding or gemmation. The process begins with the projection of the inner layer of the cell or endosporium, following Hektoen, which pushes the transparent zone and outer membrane in front of it. The bud becomes enlarged and surrounded by the same walls as surround the mother cell, and division takes place by the presence of a cross-wall which is formed by the pinching in of the cell (pi. 6, figs. 1-3, 6-7). On artificial media, on the other hand, proliferation of the fungus is brought about through sexual reproduction which is heterogamous. Two terminal cells may fuse (pi. 6, fig. 19) or two hyphae growing side by side may send out lateral cells which copulate (pi. 6, fig. 18). In either case, a spherical ascus results which may be terminal on a long filament or lateral on a short peduncle (pi. 6, fig. 33) and has a thick capsule (pi. 6, figs. 21 and 33), sometimes surrounded by a sheath as in pi. 6, fig. 35. The latter case, however, is rare and was 98 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 20 observed only three times. There are formed 8 spherical to ovoid, smooth spores which are hyaline when young and held in a gelatinous substance and when older become chamois-colored and granular, and have at maturity a diameter of 2-3 \x, varying on different media (pi. 6, figs. 38 and 33). This organism therefore agrees with that described by Gilchrist and Stokes as Blastomyces dermatitidis, but further observation has here been made on the sexual development. Cultural Descriptions The culture in this case was growing on glucose-glycerine agar, being an inoculum from an abscess on the left arm. All cultures used in the cultural determinations were taken from the above tube and grown at room temperature, approximately 22° C. Having the two stages so common with yeast-like organisms and characteristic of several members of the Endomycetales, it was thought desirable to transfer the fungus on a wide variety of media and pH range. Possessed with saccharomycetous prop- erties, on the one hand, and filamentous fungous affinities, on the other, the above method of culturing proved to be satis- factory in this work. The following media used are arranged in the order of decreas- ing hydrogen-ion concentration. Rauliri's Solution (pH I+.l). — Culture shows a thin, smooth suspension of yeast-like cells, budding and branching, varying in size from 4^-5J^ \k x 7-8^ [x, with several showing a change to filamentous formation. Richards' Solution Agar (Media consisting of Richards 7 solution with the addition of 1.5 per cent agar. pH 44)- — Growth sparse, of fine filaments. Colony V/i cm. in diameter at end of 30 days. Culture shows long hyphae projecting from edge of growth, 2^- 3 [x in diameter, with numerous budding cells. Filaments branching, with cross-walls, numerous chlamydospores, and swellings. Characteristics of the group present. Color of colony isabella to cinnamon, strongly suggestive of chamois, due to the spores and asci which are in abundance. CzapeWs Agar (pH Jf.Ji). — Color of colony white, becoming chamois with age. Growth very sparse and cottony, with much 1933) MOORE — BLASTOMYCOSIS 99 of the mycelium submerged in the agar. Colony 4 cm. in diam- eter at end of 24 days. Hyphae long and thin, lV^-2 p in diameter, with swellings approximately 4 x 12 \l, several thick- walled chlamydospores lYi x 14 y., and numerous terminal hypnospores. Several 8-spored asci seen, as well as many conidia. Malt Extract Agar (pH 5.8). — Growth slow and cultural characteristics insufficiently different to be taken into discussion. Sabouraud's Agar (pH 5.6). — Growth rapid, profuse, obtaining a diameter of l]/2 cm. at end of 30 days. Culturally the colony simulates very much that of Microsporon audouini of Ota and Langeron in the presence of several radiating ridges from the round center, the inoculum, and the several concentric rings of growth of decreasing abundance, just outside the ridges. Color of colony white when young and becoming the characteristic chamois when older. Like M . audouini, it has racquet mycelium, chains of round cells on a hypha measuring 3}^ y. in diameter, numerous chlamydospores 8 x 12 ti, terminal hypnospores 5 x 11 [l, and many conidia, characters found also in the Tricho- phytons and peculiar to Endomyces capsulatus. Unlike M. audouini, however, this organism reproduces by the formation of asci which are numerous here, measuring from 10 to 13 (x in diameter, containing 8 spores. Sabouraud's Broth (Sabouraud's medium minus the agar. pH 5.6). — Culture consists of submerged mycelium of large flakes, each measuring approximately 2 cm. in diameter at end of 24 days. Mycelium floating on surface, dry, and chamois colored, with white region, presumably the young hyphal elements. In general, growth is good. Microscopically, the culture shows long, narrow hyphae 2)4, ix in diameter, branching and inter- twining. Submerged mycelium shows almost no swellings, chlamydospores, terminal hypnospores, nor thick-walled cells, as compared with the great number found in that on the surface. The several that are present, however, show a great reduction in size and form from the exposed, the measurements of which are similar to those on agar. Potato-dextrose Agar (pH 5.6). — Growth profuse and cottony, covering the surface of the agar completely. Diameter of colony [Vol. 20 100 ANNALS OF THE MISSOURI BOTANICAL GARDEN l]/2 cm. after 24 days. Color cinnamon, with colony showing concentric circles of color alternating with white, and the cinna- mon very pronounced, due perhaps to the medium constituents. Hyphae 3 [x in diameter, with numerous, thick-walled cells 7^ [x in diameter, budding cells, swellings 5 x 12 [a, and chlamydospores varying in size from 4-7 x 9-14 y,. Asci numerous, measuring approximately 13 [l in diameter. Corn-meal Agar (product of Digestive Ferments Co. pH 6.0).— Growth poor, colony being V/i cm. in diameter at end of 24 days. Color white. Growth around inoculum loose and cottony. Hyphae short, thick-walled, 2]^ V- in diameter, with numerous budding cells approximately 7 [x in diameter. Chlamydospores numerous, 7 x 13 ix, terminal hypnospores several, 5 x 12 [x; a few asci seen, 11 (x in diameter. Conidia abundant, 5 ix in diameter. June-beetle Agar (medium consisting of a 4 V er cen t extract of June beetles, Lachnosterna fusca, plus 1.5 per cent agar, sterilized at 20 pounds pressure for 20 minutes, with a final pH 6.1). — Growth of loose, flat, cottony mycelium, forming concentric circles of decreasing abundance until a ring of fine filaments surrounds the culture. Colony h l /2 cm. in diameter at end of 24 days. Hyphae 2-23/2 tx in diameter, with many conidia 5 jx in diameter. Asci 12-13 ix in diameter, thick-walled, enclosed in a sheath. Abundance of racquet mycelium. June-beetle Dextrose Agar (above medium plus 2 per cent dextrose). Growth fair, attaining a diameter of 3 cm. at end of 30 days. Colony bright chamois in color, cerebriform, and cottony. Many conidia, 4^-5 \i in diameter. Hyphae 2^-3 (x in diameter and fairly short. Chlamydospores 8 x 16 ;x and numerous, as well as terminal hypnospores 5x12 [l. Asci round, 12-14 \i in diameter. Lactose Agar (product of Digestive Ferments Co., lactose broth plus 1.5 per cent agar. pH 6.8). — Growth good, reaching a diam- eter of G cm. at end of 24 days. Colony chamois-color, profuse and cottony, with a region of very fine mycelium surrounding it. Hyphae 3^ [x in diameter with numerous conidia 5 y. in diameter, budding oil. Many thick-walled resting cells 7 n in diameter. Characteristic racquet mycelium, chlamydospores, terminal hypnospores, with properties similar to those on Sabouraud's agar. 1933] MOORE — BLASTOMYCOSIS 101 Lactose Broth (product of Digestive Ferments Co. pH 6.8), Growth good, large white flakes being formed in the solution which later become intertwined, forming a mat of mycelium 7 [x in diameter. Hyphae slightly reduced, 3 u, in diameter, 1 Vo u in the younger filaments. Preponderance of budding cells 7 (x in di ameter, with thick- walled chlamydospores, asci, and terminal hyp but reduced in size as compared with the growth agar. E osine-methylene-blue Agar (agar used as one of a routine, prod- uct of Digestive Ferments Co. pH 7.0). — Growth good, with a diameter of 534 cm. at end of 24 days. Culture compact, due to the hyphae having absorbed the stain from the substrate and turning the mycelium pink. Colony appears powdery with age. Hyphae characteristic, with swellings, 3 n in diameter. Many conidia, 5 p in diameter, and hypnospores with several chlamy- dospores. Glycerine Agar (nutrient agar as prepared by the Digestive Fer- ments Co. plus 6 per cent glycerine, Merck C. P. pH 7.1). — Growth fair, having a diameter of 5 cm. at end of 24 days. Culture shows a crinkled, moist region of budding yeast-like cells and a dry filamentous, cottony, chamois-colored region which has changed to the mycelial form characteristic on agar. Filamentous hy- phae 3}4 [i in diameter, characteristic swellings being present which are slightly larger than those found on lactose agar. Racquet mycelium also present. Nutrient Agar (product of Digestive Ferments Co. pH 7.2). Growth rapid, covering a region 7 cm. in diameter at end of 30 days. Colony filamentous, cottony, brown, with concentric rings of growth, the outermost being white. Hyphae 234 v. in diam- eter. Growth similar to that on Sabouraud's agar micro- scopically, with numerous conidia 5 ^ in diameter and asci 13 tx in diameter. Nutrient Broth (pH 7.2). — Culture forms a mat of intertwining mycelium of long hyphae 2-234 v- in diameter, with swellings, asci, and chlamydospores. Very few conidia. Terminal hyp- nospores several, but reduced in size, 4 x 9 (x. Endo's Agar (product of Digestive Ferments Co. pH 7.5). — Growth fair, colony having a diameter of 334 cm. at end of 30 102 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 20 days- Culture shows radiating ridges from center of inoculum, with growth becoming flat due to the stain in the medium which is absorbed by the hyphae, as in the case of the eosine-methylene- blue agar, giving the mycelium a pink color. Microscopically, the hyphae have a diameter of 2^ p. Numerous conidia 5 \l in diameter. Culture otherwise similar to that on eosine- methylene-blue agar. Gelatine {nutrient agar plus 1.5 per cent gelatine). — Slow lique- faction beginning after 30 days. Culturally, the fungus is very characteristic of the organism of blastomycosis in that it passes through the three typical stages: the moist, yeast-like stage with a flat growth; the prickly culture with the colonies simulating greatly small burrs (coremia); and the final, cottony growth present on agar after extended growth. Discussion As stated in the introduction of this paper, it would seem that the syndrome-complex, commonly known as blastomycosis, has an innumerable list of etiological factors, each causing a condition so much like the other that clinicians have grouped them under one head. However, should one encounter any of these in a clinic one would find that therapeutic measures are so vastly different, varying with the organism, that a direct and accurate knowledge of the causative agent in each particular patient is absolutely essential. In the past, medical men, not particularly trained in mycologi- cal taxonomy, were inclined to class together all fungi presenting ascomycetous characters under one name, Blastomyces. So great is the confusion to-day that it is necessary to pick out these pathogenic fungi and classify each one separately. The organism isolated in the first case was termed Blastomyces demiatitidis by Gilchrist in 1894 because of its budding properties in the lesion. In a case of dermatitis reported by Gilchrist and Stokes ('96) the organism, which was evidently of the type termed Blastomyces, was called an Oidium. In a following paper (Gil- christ and Stokes, '98), it was made known that the organism described in the previous paper was called an Oidium because it did not ferment glucose, saccharose, or lactose, and although 1933] MOORE — BLASTOMYCOSIS 103 developing by gemmation or budding in the tissues, human and animal, developed mycelia with the formation of conidia upon artificial media. Ricketts ('01) made an extensive study of the organism, distinguishing it from several of the yeasts but failing to consider several of the yeast-like fungi, and proposed definitely the name Oidium for the genus of the Gilchrist fungus. After this work, several terms were applied to the disease. Busse ('94) described his case a short time after Gilchrist reported his and he named the organism Saccharomyces hominis. Vuillemin in a later publication assigned the organism to the genus Cryptococcus and called it C. gilchristi. However, he failed to make a careful study of the organism culturally on artificial media, and a classification which places a great emphasis on the yeast-like appearance of the fungus in lesions is not exactly justifiable. Brumpt ('27) places the organism in the genus Mycoderma, calling it M. dermatitis. This terminology, however, is synony- mous with Oidium, and in that case is likewise useless. For a great number of years, no great work was done to establish definitely the position of Gilchrist's organism, and the name Blastomyces as created by him still held sway. The term pre- sents a lot of difficulties. In the first place, the Blastomycetes, according to Buschke, are that group which develops through budding, provided a mycelium is formed on agar, while to the group of Blastomycetes, as Naegeli names budding, would belong the genera Endomyces, Saccharomyces, Cryptococcus, Monilia, and Oidium. Now the question arises as to what the actual meaning of the word blastomycete is. According to Vuillemin ('01), it does not designate a natural group, a botanical family based on genealogical affinities. There is in existence a genus Blastomyces, but these organisms are not budding fungi in the sense of Buschke. They are filamentous fungi whose spore-bearing elements, whether terminal, lateral, or intercalary, can be isolated by disarticulation, following Costantin and Rolland (quoted by Vuillemin). Frank (quoted by Vuillemin) established the Blastomycetes as an order to include such fungi as the beer yeasts whose elements are iso- lated by budding and not by disarticulation. In this respect, by virtue of the law of priority, it would seem that the name Blas- tomycetes, as designated by Frank, should remain. However, [Vol. 20 104 ANNALS OF THE MISSOURI BOTANICAL GARDEN if by general agreement the name of a genus could replace that of an order, then, according to the rules of nomenclature, the genus of Costantin and Rolland is legal, and the name as designated here is not legitimate by reason of the lack of distinct characters which have no generic value. Castellani recently proposed a new classification of yeast-like or budding fungi based on the presence or absence of ascospores, which includes families of both the Ascomycetes and Fungi Im- perfecti. 1. Saccharomycetaceae : budding cells, asci, and ascospores, but no mycelium in culture. 2. Endomycetaceae : budding cells, asci, and ascospores, with mycelium in culture. 3. Cryptococcaceae : budding cells (blastospores), no asci and no mycelium in culture. 4. Oosporaceae : budding cells, no asci, but mycelium in culture. In addition to this family classification, he created a new genus which he calls Blastomycoides, to which he assigns three species, and places it in the family Oosporaceae: 1. Blastomycoides der- matitidis, synonym Blastomyces dermatitidis Gilchrist and Stokes; 2. Blastomycoides immitis, synonym Coccidioides immitis Rixford and Gilchrist; 3. Blastomycoides tulanensis Castellani. He de- fines the genus Blastomycoides as: "Oosporaceae appearing in the lesions as large roundish cells from eight to twenty microns in diameter, or larger, with the protoplasm containing a number of well-marked granules or spherules, and with a membrane showing a well-defined double contour; in dextrose agar cultures a large amount of mycelium is present." He bases further differentiation of the three species on their cultural differences when grown on mannitol, lactose, glucose, and galactose agar. The second species that he names, Blastomycoides immitis, has already been discussed and classified by the author in a previous paper (M. Moore, '32). The author has made no pretext of studying the third species, so that nothing can be said about that. The first species, however, Blastomycoides dermatitidis, is alto- gether misplaced, simply because there are asci present in the mycelium in culture. This of course would refer the genus to the family Endomycetaceae, in which group the writer definitely establishes the organism. 1933] MOORE — BLASTOMYCOSIS 105 Observations on the growth, development, reproduction, and further evolution of the fungus show that there are budding cells in the lesions, mycelium formed on agar with an intermediate stage showing the change from the yeast-like to the filamentous form. In accordance with this, Mellon ('24, '26, J 26a) has recorded the fact that asci do occur particularly "in the so-called secondary colonies of the cultures which also contained 'dau- ernzellen' and pigmented oidia." The author wishes to affirm Mellon's findings as to the presence of asci, but suggests that these structures are present in the third stage, whereas the second step would consist of the intermediary forms which have an appearance very much like greatly enlarged oidia. Furthermore, it would seem that Mellon's description was indefinite, inasmuch as he refers to an ascus as an ascospore, and oil droplets are suggested by him as being chromatin indicators and forerunners of the future spores. Such factors as these are very important in the taxonomy of this type of fungi and should not be dealt with so promiscu- ously. Furthermore, Mellon has not paid much attention to the fact that no matter how old the lesion may be the blastomycosis organism found there does not change from its yeast-like, budding growth until it has been transferred to artificial media, where the change is an adaptation to the mode of life it must lead; in other words, the change from active parasitism to one of saprophytism. It is to be understood, however, that a change such as suggested here will not necessarily reduce its viability, at least for the time being. A study of the evolution of the organism has repeatedly shown, in several hanging-drop cultures, that reproduction is hetero- gamous, as given in the description, with the final formation in the series of a large eight-spored ascus. Mellon in his papers consistently shows a four-spored ascus. It would seem, therefore, that he either has an organism unlike the one here described for the blastomycosis parasite, or else he has taken for granted as spores the four oil droplets which may and often have been found to occur on a mature eight-spored ascus, as was evidenced by the author on another ascomycete, Endomyces capsulatus var. isabellinus Moore, which was described in a case in another paper (McBride and Thompson, '33) and also in Endomyces [Vol. 20 106 ANNALS OF THE MISSOURI BOTANICAL GARDEN capsulatus Rewbridge, Dodge and Ayers ('29). Furthermore, it is quite possible that Mellon has observed the ascus just previous to the division of the nuclei, in the formation of the eight-spored ascus. This latter statement is only a conjecture on the part of the writer, but in any case the cultural descriptions do not agree with those given by the early investigators and with which the organism here described does agree. In view of such criteria, it would seem that the organism formerly described as Blastomyces dermatitidis Gilchrist 1894, is not strictly a member of that genus, the name of which, on account of its etymological derivation, is essentially a misnomer. Because of its morphological characteristics, Blastomyces derma- titidis does not present those affinities entirely but simply as one phase of its life cycle. However, particularly because of its asco- mycetous attributes, it should belong to the class Ascomycetes, order Kndomycetales, family Endomycetaceae, and because of its similarity in morphology and reproduction (perfect stage) to that of Endomyces capsulatus and its variety, it should belong to the genus Endomyces. If taxonomic position in this family and genus be dependent on the number of spores in the ascus, it would seem, according to Whitman ('13), that this organism should belong in the genus Oleina. However, the genus Endomy- ces contains a number of pathogenic species with eight-spored asci, whereas Oleina has no pathogenic species, and until a classifica- tion better than the one now in existence be established, dermati- tidis should be placed with Endomyces. From the above statement it would appear that the organism should now be known as: Endomyces dermatitidis (Gilchrist 1894), M. Moore, n. comb. Mycelium in lesions of budding yeast-like cells 7-12 \i in diam- eter and sometimes as much as 20 [x in length, occurring singly, in groups of two's, three's, or four's. Growth on agar of isodia- metric cells 2-2)^ \l in diameter on acid media and 3-4 [i in diam- eter on slightly alkaline media. Hyphae septate, with conidia pyriform or round, pedunculate or sessile, 5 (jl in diameter. Racquet mycelium present, 5-6 \l in diameter at swollen portion and 3-3j/£ [l in diameter at narrow portion. Chlamydospores terminal or lateral or intercalary, ^Yr^Vi x 12-15 [jl, or sometimes 1933] MOORE — BLASTOMYCOSIS 107 round, 7 p. in diameter. Copulation heterogamous, asci spherical 8-13 (x in diameter, with 8 spherical to ovoid, smooth, hyaline to light chamois-colored spores 2-3 p in diameter, at maturity Colony white in color, becoming cinnamon to brown with age. Summary 1. The history of blastomycosis is given, with a review of the early work on yeast-like, fungous pathogenicity, and a report of the first case published. 2. The etiology and clinical manifestations represent a number of conditions due to several yeast or yeast-like organisms: Sac- charomyces, Oidium, Monilia, Endomyces, Cryptococcus, and Coccidioides, which have been placed in one category to constitute the agents responsible for the syndrome-complex, blastomycosis. 3. The disease is shown to simulate several conditions, in which cases the diagnosis must be arrived at through the isolation of the organism and the application of Koch's postulates. 4. Immunological reactions and therapeutic measures are as yet indefinite as to specific results, although beneficial results have been reported by the use of iodides. 5. A case of blastomycosis of the arm and hand is reported. 6. There is a description of the organism, culturally and mor- phologically, showing its relationship to the class Ascomycetes. 7. The fungus is definitely established as Endomyces dermati- tidis of the family Endomycetaceae. Acknowledgments The author wishes to express his sincere gratitude to the following: Dr. Carroll W. Dodge, Professor of Botany in the Henry Shaw School of Botany of Washington University, for his interest, criticisms, and helpful suggestions; Dr. George T. Moore, Director of the Missouri Botanical Garden, for the courtesies extended; Dr. Martin F. Engman, Dermatologist to the Wash- ington University Hospitals, for the use of the data with respect to the case reported; and Miss Nell C. Horner, librarian of the Missouri Botanical Garden, for her assistance. 108 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 20 Bibliography Agostini, A. ('31). On Blastomycoides lanuginosus Castellani. Jour. Trop. Med. &Hyg. 34:287-288. 1931. Basgal, W. ('31). Contribuigao ao estudo das blastomycoses pulmonares. Doctor- ate thesis in medicine. Rio de Janeiro, 1931. Bassoe, P. ('00). Report of a case of disseminated blastomycosis of the lungs, lumbar vertebrae and subcutaneous tissues. Chicago Path. Soc, Trans. 6: 380. 1900. Benedek, T. ('28). Bemerkungen zum Zuchtungsverfahren des Schizosaccharo- myces hominis Benedek, 1927. I Mitteilung. Die Primarkultur. Derm. Wochenschr. 87: 1203-1214. 1928. , and 11. Friihwald ('28). Clinical picture, mycology and serum diagnosis of schizosaccharomycosis; 2 cases. Ibid., 1566-1577. 1928. Blanchard, R., E. Swartz, et J. Binot ('03). Sur une blastomycose intrap6riton6ale. Arch, de Parasitol. 7: 489-507. 1903. Bigot, A., et H. Velu ('25). I so lenient rapide de CryptoCOCCUl mirandei en culture pure. Soc. Path. Exot., Bull. 18: 127-129. 1925. , , ('25a). Etude biologique de Cryptococeus mirandei agent de la blastomycose des voies lacrymales de P&ne. Ibid., 231-235. 1925. , , ('25b). Contribution i\ Petude des blastomycoses animates. Rev. Path. Comp. et Ilyg. G6n. 25: 280, 281, 283. 1925. Borzone, R. A. ('29). I'n caso de blastomicosis en Santa F6 y ensaya de revisi6n de las blastomicosis americanas. Soc. Scient. Santa Fe, An. 1: 58-62. 1929. Bowen, J., and S. B. Wolbach (W>). A case of blastomycosis; the results of culture and animal experiments. Jour. Med. Res. 10: 1 (17-177. 1906. Brown, P. K., and W. T. Cummins ('15). I. A differential study of coccidioidal granuloma and blastomycosis. II. Report of two additional cases of cocci- dioidal disease. Arch. Int. Med. 15: 608-627. 1915. Brumpt, E. ('27). Prdcis de parasitologic pp. 1213, 1383, MassonetCie. Paris, 1927. Burkhead, C. E. ('22). Oidiomycosis, including one case of coccidioidal granuloma and one of cutaneous blastomycosis. Kan. Med. Soc, Jour. 22: 101. 1922. Buschke, A. (9S). Ueber Ilautblastomykosc Deutsch. Derm. Gesell., Verhandl. 6: 181-222. 1898. , und A. Joseph ('28). Blastomykose (Ascomykose). In Jadassohn, Hand- hue h der Ilaut- und Geschleehtskrankheiten 11: S25-925. 1928. Busse, < >. ('94). Ueber parasitaire Zelleinschlusse und ihre Ziichtung, Centralbl. f. Bakt. Orig. 16: 175-180. 1894. , ('95). Ueber Saccharomycosis hominis. Virch. Arch. 140: 23-46. 1895. Castellani, A. ('25). Observations on some diseases of Central America. (Blas- tomycosis in man in Central America.) Jour. Trop. Med. & Hyg. 28: 1-14. 1925. , ('25a). Notes on three new yeast-like organisms and a new bacillus, with remarks on the clinical conditions from which they have been isolated; furun- culosis blastomycetica, macroglossia blastomycetica, stomatitis cryptococco- baeillaris. Ibid., 217-223. 1925. , ('28). Fungi and fungous diseases. Am. Med. Assoc Chicago, 1928. , ('28a). Notes on blastomycosis; its etiology and clinical varieties. Roy. Soc. Med., Sect. Trop. Dis. & Parasitol., Proc 21: 447-462. 1928. 1933] MOORE — BLASTOMYCOSIS 109 Castellani, A., ('28b). Blastomycosis and some other conditions due to yeast-like fungi (budding fungi). Am. Jour. Trop. Med. 8: 379-422. 1928. , ('29). Mannitol agar in the differentiation of the fungi of type Blastomy- ces. Soc. Exp. Biol. & Med., Proc. 26: 544. 1929. Chatenewer ('28). Material zur experimentellen Blastomykose des Kaninchens. Derm. Wochenschr. 87: 1649. 1928. Chiari, H. ('30). Zur Pathologie und Histologic der generalisierten Torulose (Blastomykose). Arch. f. Derm. u. Syph. 162: 422-441. 1930. Chyurlia, N. ('26). Notes on a case of bronchomycosis. Jour. Trop. Med. & Hyg. 29: 145-146. 1926. Cleary, J. H. ('04). A case of generalized blastomycosis. Chicago Path. Soc, Trans. 6: 105-113. 1904. Cleland, J. B. ('27). A case of systemic blastomycosis with the formation of a myxomatous-looking tumor-like mass. Med. Jour. Australia 14: 337-340. 1927. Cole, W. H. ('24). Systemic blastomycosis. Ann. Surg. 80: 124-134. 1924. Copelli, M. ('13). A case of blastomycosis. Jour. Cut. Dis. 31: 51-52. 1913. Corselli, G., und B. Frisco ('95). Pathogene Blastomyceten beim Menschen. Beitrage zur Aetiologie der bosartigen Geschwiilste. Centralbl. f. Bakt. 18: 368-373. 1895. Coupal, J. F. ('24). Diagnosis and treatment of certain disease entities. Report of six cases of blastomycosis. Internat. Clin. 4: 1-14. 1924. Curtis, F. ('96). Contribution k Petude de la saccharomycose humaine. Inst. Past., Ann. 10: 449-468. 1896. Davis, B. F. ('11). The immunological reactions of oidiomycosis (blastomycosis) in the guinea-pig. Jour. Inf. Dis. 8: 190. 1911. , ('22). Blastomycosis: Clinical pathology and therapeusis. Minn. Med. 5:311-315. 1922. Davis, C. N. ('06). A case of blastomycetic dermatitis. Jour. Cut. & Vener. Dis. 24: 90. 1906. Dennis, F. L. ('18). Blastomycosis of the upper respiratory tract with a report of a case primary in the larynx. Ann. Otol., Rhin. & Laryng. 27: 571. 1918. Desjardins, A. U. ('25). Roentgenotherapy and diathermy in blastomycosis. Am. Jour. Roentgenol. 14: 14-16. 1925. Dowling, G. B., and R. R. Elworthy ('25). A case of blastomycetic dermatitis (Gilchrist). Roy. Soc. Med., Proc. 19: 4-10. 1925. Downing, E. D. ('18). A case of blastomycosis with laryngeal involvement. Am. Med. Assoc, Jour. 70: 85-86. 1918. Eisendrath, D. N., and O. S. Ormsby ('05). A case of systemic blastomycosis in the sputum. Ibid. 45: 1045. 1905. Engelhardt, W. ('24). Ein Beitrag zur Aetiologie oberflachlicher Hautblastomyko- sen und Hautsoormykosen. Arch. f. Derm. u. Syphil. 146: 313-322. 1924. Evans, N. ('03). A clinical report of a case of blastomycosis of the skin from acci- dental inoculation. Am. Med. Assoc, Jour. 40: 1772-1775. 1903. , ('09). Coccidioidal granuloma and blastomycosis in the central nervous system. Jour. Inf. Dis. 6: 523-526. 1909. Fabry, J. ('25). Superficial erosive blastomycosis. Derm. Wochenschr. 81: 1071- 1075. 1925. , ('27). Vbev akneformige blastomycosis cutis. Ibid. 84: 824-827. 1927. 110 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 20 Ferguson, A. S. ('28). Blastomycosis of eye and face secondary to lung infection. 442-443. 1928. da Fonieca, O. ('22). Sobre as agentes das blastomycoses europeas. Cyclosexuadoe possigao systematico do levedo de Hudelo. Brasil-Med. 36: 101-102. 1922. , ('28). Ensaya de revisi6n de las blastomicosis sudamericanos. Inst. Clin. Quirurg., Hoi. 4: 469-502. 1928. , et A. E. de Area Lefio ('28). Dermatite blastomycosique. Soc. Biol., Compt. Rend. 98: 622-623. 1928. Fontaine, B. W., M. Haase, and R. H. Mitchell ('09). Systemic blastomycosis. Arch. Int. Med. 4: 101-117. 1909. Forgues, J. B. C. ('13). Contribution a l'dtude des exoasc<5es pathog&ies. Thfese de Bordeaux, 100 pp. 1913. Foulerion, A. ('00). On the pathogenic action of blastomycetes. Jour. Path. & Bact, 6: 37-63. 1900. Freeman, W., and F. D. Weidman ('23). Cystic blastomycosis of cerebral gray matter caused by Torula histolytica Stoddard and Cutler. Arch. Neurol. & Psychiat. 9: 589 603. 1923. Froilano de Mello, et A. Rodrigues ('29). Sur un cas de blastomycose k placards multiples v6g6tants verruqueux ou pustulo-ulceres. Soc. Path. Exot., Bull. 22: 142-147. 1929. Garr, C. C. ('25). Systemic blastomycosis. Surg. Gyn. & Obs. 41 : 490-492. 1925. Gdspdr, I. ('29). Blastomycotic meningoencephalitis. Arch. Neurol. & Psychiat, 22: 475-486. 1929. Gilchrist, T. C. ('96). A case of blastomycetic dermatitis in man. Johns Hopkins Hosp., Repts. 1: 269-283. 1896. , ('02). Blastomycetic dermatitis in the negro. Brit. Med. Jour. 2: 1321- 1328. 1902. , and W. R. Stokes ('96). The presence of an Oidium in the tissues of a case of pseudo-lupus vulgaris. Johns Hopkins Hosp., Bull. 7: 129-133. 1896. f , ('98). A case of pseudo-lupus-vulgaris caused by a blastomyces. Jour. Exp. Med. N. Y. 3: 53-78. 1898. Graves, M. L. ('22). Systemic blastomycosis. Am. Jour. Trop. Med. 2: 123-132. 1922. Greenfield, J. G. ('24). Blastomycosis of nervous system. Med. Sci. 10: 267-273. 1924. Grschebin, S. ('27). Ein Fall von tiefer primiirer Blastomykosis der Haut (Busse- Buschke). Derm. Wochenschr. 85: 1019-1055. 1927. ■ , ('28). Deep primary blastomycosis of the skin. Urol. & Cutan. Rev. 32:453 1")7. 1928. , und L. N. Maschkilleisson ('26). Beitrage zur Lehre von der pathologi- schcu Anatomic der Gilchristchen Hautblastomykose. Derm. Wocher 82: 811-818. 1926. Haase, M., E. R. Hall, and C. H. Marshall ('22). Local blastomycosis, report of a case. Am. Med. Assoc, Jour. 79: 820-822. 1922. Hagiwara, S. ('22). Uber Blastomycosis cutis. Jap. Zeitschr. Derm. Urol. 22: 941-980. 1922. Hamburger, W. W. ('07). A comparative study of four strains of organism isolated from four cases of generalized blastomyeosis. Jour. Inf. Dis. 4: 201-209. 1907. Hamilton, C. M. ('26). Blastomycosis. South. Med. Jour. 19: 431-435. 1926. 1933] MOORE — BLASTOMYCOSIS 111 Harter, A. ('09). De la blastomycose humaine. These Fac. M6d. Nancy 8: 222. 1909. Hashimoto, T. ('22). tTber Blastomycosis cutis. Jap. Zeitschr. Derm. Urol. 22: 1-34. 1922. Hedge, H. M. ('28). The use of carbon dioxide snow in treating blastomycosis. Am. Med. Assoc, Jour. 90: 1367-1369. 1928. Hektoen, L. ('99). The organism in a case of blastomycetic dermatitis. Jour. Exp. Med. 4: 261-278. 1899. f ('07). Systemic blastomycosis and coccidioidal granuloma. Am. Med. Assoc, Jour. 49: 1071-1077. 1907. Herrick, J. B. ('07). Generalized blastomycosis. Ibid. 328. 1907. Hessler, R. ( , 99). Blastomycetic dermatitis. Ibid. 32: 760. 1899. Hicks, J. A. B., and F. R. Chopping ('24). Case of perionychia due to a blastomyces. Lancet 206 1 : 128. 1924. Hill, H. P., and E. C. Dickson ('14). Report of a case of systemic blastomycosis. Calif. State Jour. Med. 12: 120. 1914. Howes, W. B., and P. F. Morse ('21). Report of two cases of blastomycosis. Boston Med. & Surg. Jour. 185: 315-317. 1921. Hudelo, Rubens-Duval, et Laederich ('06). fitude d'un cas de blastomycose k foyers multiples. Soc M6d. H6p. Paris, Bull, et M6m. 23: 723-734. 1906. Hufschmitt, G., A. Sartory, R. Sartory, et J. Meyer ('31). Un cas de blastomycose cutan6e k foyers multiples. Ann. Dermatol. 7: 850-876. 1931. Hurley, T. D. ('16). Heart lesion in blastomycosis. Jour. Med. Res. 33: 499-502. 1916. Hyde, J. N., L. Hektoen, and A. D. Bevan ('99). A contribution to the study of blastomycetic dermatitis. Brit. Jour. Derm. 11: 261-276. 1899. Irons, E. E., and E. A. Graham ('06). Report of a case with miliary and ulcerative blastomycosis of the lungs. Miliary blastomycosis of the spleen and multiple superficial and deep abscesses. Jour. Inf. Dis. 3: 666-682. 1906. Jackson, C. ('26). Blastomycosis of the larynx. Arch. Otolaryng. 3: 99-107. 1926. Jackson, R. H. ('26). Surgical treatment of certain massive blastomycetic skin lesions. Am. Jour. Surg. 1: 185-187. 1926. Jacobson, H. P., J. F. Schamberg, and H. Morrow ('32). Fungous diseases. A clinico-mycological text. pp. 149-181. Charles C. Thomas Co., Springfield, Illinois. 1932. Jeaume, G., et M. Dekester ('25). Isolement de l'agent pathogene de la blastomy- cose des voies lacrymales. Soc. Path. Exot., Bull. 18: 124-127. 1925. Jona, G. ('97). Die Schutzmittel des Organismus gegen Blastomyceten. Centralbl. f. Bakt. 21: 147-150. 1897. LeCount, E. R., and J. Myers C05). Systemic blastomycosis. Final report of the case described by Eisendrath and Ormsby in 1900. Jour. Inf. Dis. 4: 187-200. 1905. Legendre, J. ('27). A propos de la dermatite blastomycosique ch61oidienne. Soc. Path. Exot., Bull. 20: 323. 1927. Lewis, D. ('17). Blastomycosis and sporotrichosis. Surg. Clinics, Chicago 1: 1125. 1917. MacLeod, J. M. H. ('30). Some skin affections due to yeast-like fungi. Brit. Med. Jour. 1930: 1119-1123. 1930. 112 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 20 Maffuci, A., unci L. Sirleo ('98). Ueber die Blastomyceten als Infektionserreger bei bflaartigen Tumoren. Zeitschr. f. Ilyg, 27: 1-30. 1898. Manor, G. D., and It. W. Hammack ('30). Systemic blastomycosis. Calif. & Went. Med. 32: 87-90. 1930. Massey, A. Y. ('16). Blastomycosis (?) in Central Africa. Jour. Trop. Med. & Hyg. 19: 79. 1916. Mazza, 8., y F. Nino ('28). Notas sobre blastomicosia de las vias respiratorias. Reunion Soc. Argentina Patol. Reg. Norte en Santiago del Estero 4: 545-548. 1928. , , II. Quintana, y V. Bernasconi ('30). Blastomicosis grave geueralizada por Monilia n. sp. Reuni6n Soc. Argentina Patol. Reg. Norte en Salta 6: ISO 214. 1930. , y F. Canevari ('29). Ulceras blastomicosicns de la lengua. Reuni6n Soc. Argentina Patol. Reg. Norte 5: 220-230. 1929. , F. L. Nino, y P. Nicolini ('29). Blastomicosis de la mucosa labiogeniana. Ibid. 231-239. 1929. , , y A. Egiies ('29). Perionixis blastomieetiea por Monilia (n. sp.). Ibid. 284-288. 1929. , L. Stabile de Nucci, y E. J. Canal Feij6o (Santiago del Estero) ('29). Blastomicosis cutdnea de forma lenta por criptococa (n. sp.). Ibid. 293-308. 1929. , y B. Palamedi ('32). Caso mortal de blastomicosis cut&neo mucosa. Reunion Soc. Argentina Patol. Reg. Norte en Tucuman 7: 424-407. 1932. McKee, S. II. ('20). Blastomycosis of the cornea, with review of reported cases of blastomycosis of the eye. Internat. Clin. 3: 50-57. 1926. Meckel, M. ('27). Weitere Mitteilungen liber erosive Blastomykosen. Derm, Wochensehr. 84: 817-824. 1927. Medlar, K. M. ('27). Pulmonary blastomycosis; its similarity to tuberculosis. Am. Jour. Path. 3: 305-314. 1927. Mellon, R. R. ('24). Observations on an ascospore stage for the parasites of blas- tomycosis hominis. Exp. Biol. & Med., Proc. 22: 09. 1924. , ('26). Studies in microbic heredity. VI. The infective and taxonomic significance of a newly described ascospore stage for the fungi of blastomycosis. Jour. Bact. 11: 229-252. 1926. , ('26a). Studies in microbic heredity. VII. Observations on the genetic origin of the several types of fungi found in t he lesions of blastomycosis hominis. Ibid. 419-432. 1926. Michelson, I. D. ('28). Blastomycosis; pathologic and bacteriologic study. Am. Med. Assoc, Jour. 91: 1871-1S76. 1928. Miller, J. E. ('25). Yeast-cell formation in man. U. S. Navy Med. Bull. 23: 229- 235. 1925. Miller, W. S. ('27). The reticulum of the lung: Its similarity in blastomycosis to that in tuberculosis. Am. Jour. Path. 3: 315-320. 1927. Montel, R., and It. Pons ('26). Dermatite blastomycosique ch61oidienne. Soc. Path. Exot., Bull. 19: 876-880. 1926. Montgomery, F. H. ('03). A case of cutaneous blastomycosis followed by laryngeal and systemic tuberculosis. Death; autopsy. Jour. Cut. Dis. 21 : 19-22. 1903. , and (). S. Ormsby ('08). Systemic blastomycosis: Its etiologic, pathologic and clinical features as established by a critical survey and summary of twenty- 1933] MOORE — BLASTOMYCOSIS 113 two cases, seven previously unpublished. The relation of blastomycosis to coccidioidal granuloma. Arch. Int. Med. 2: 1-41. 1908. Montpellier, J., et A. Catanei ('26) . Blastomycose de Pavant-bras chez une femme indigene d'Alger. Soc. Path. Exot., Bull. 19: 586-592. 1926. Moore, J. T. ('20). Blastomycosis. Report of a case dying from abscess of brain. Surg., Gyn. & Obs. 31: 590-594. 1920. Moore, M. ('32). Coccidioidal granuloma: A classification of the causative agent, Coccidioides immitis. Mo. Bot. Gard., Ann. 19: 397-428. 1932. [- ], MacBryde, C. M., and E. J. Thompson. ('33). Meningitis and dermatitis caused by a new variety of blastomycete (endomycete). Arch, of Derm. & Syphil. 27: 49-69. 1933. Morris, R. T. ('13). A case of systemic blastomycosis. Am. Med. Assoc, Jour. 61: 2043-2044. 1913. Nesczadimenko, A. ('99). Zur Pathogenese der Blastomyceten. Centralbl. f. Bakt. 25: 55-58. 1899. Neumayer, J. ('91) . Untersuchungen liber die Wirkungen der verschiedenen Hefearten, welche bei der Bereitung weingustiger Getranke vorkommen auf den thierischen und menschlichen Organismus. Arch. f. Hyg. 12: 1-60. 1891. New, G. B. fl7). Blastomycosis of the tongue. Am. Med. Assoc, Jour. 68: 186. 1917. , ('28). Blastomycosis of the larynx. Ann. Otol., Rhin. & Laryng. 37: 240-250. 1928. Nieberle, N. ('27). Blastomycosis of skin in pig. Virch. Arch. f. path. Anat. 263: 16-24. 1927. Nino, F. L. ('29). Ulceraci6n blastomic^tica cut&neomucosa del labio inferior (Consideraciones acerca de su diagn<5stica etiol6gico). Reuni6n Soc. Argentina Patol. Reg. Norte 5: 213-225. 1929. , ('29a). Onixis y perionixis de origen blastomic6sico (Estudio clinico y micol6gico). Ibid. 270-282. 1929. , ('30). Blastomicosis humano generalizada por Criptococo (n. sp.). Reuni6n Soc. Patol. Reg. Norte en Salta 6: 117-167. 1930. , y J. Fernandez ('29). Nueva observacion de perionixis per Monilia periungualis. Reuni6n Soc. Patol. Reg. Norte 5: 282-283. 1929. y m. Palant ('30). Nuevas observaciones de onixis y peri- onixis de origen blastomicosico. Reuni6n Soc. Argentina Patol. Reg. Norte en Salta 6: 35-99. 1930. Ormsby, O. S. ('21). Blastomycosis. A practical treatise on diseases of the skin. Lea & Febiger, Philadelphia & New York. 1921. , and H. M. Miller ('03). Report of a case of systemic blastomycosis with multiple cutaneous and subcutaneous lesions. Jour. Cut. Dis. 21: 121-136. 1903. Ota, M. ('24). Essai de classification des blastomycoses pathogdnes. Ann. Para- sitol. 2: 34-61. 1924. Otis, F. J., and N. Evans ('03). Morphology and biology of the parasite from a case of systemic blastomycosis. Am. Med. Assoc, Jour. 41: 1075-1082. 1903. Panja, G. ('25). A case of generalized blastomycosis. Ind. Med. Gaz. 60: 475-476. 1925. Parker, C. A. ('23). Actinomycosis and blastomycosis of the spine. Jour. Bone & Joint Surg. 5: 759-777. 1923. 114 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 20 Parmenter, F. J., and B. T. Simpson ('19). A case of blastomycosis involving the prostate and seminal vesicles. Jour. Urol. 3: 449. 1919. Rabinowitsch, L. ('96). Untersuchungen liber pathogene Hefearten. Zeitschr. f. Hyg. u. Infektionskrank. 21: 11-24. 1896. Raum, J. ( ? 91). Zur Morphologic urui Biologic der Sprosspilze. Ibid. 10: 1-50. 1891, Reed, P. A. ('26). Systcrnic blastomycosis. Neb. Med. Jour. 11: 257-260. 1926. Rewbridge, A. G., C. W. Dodge, and T. T. Ayers ('29). A case of meningitis due to Endornyces capsulatus (new species). Am. Jour. Path. 5: 349-364. 1929. Rhamy, B. W. ('26). Blastomycosis of the bladder. Am. Med. Assoc, Jour. 87: 405-406. 1926. Richter, W. ('28). Beit rage zur Hefepilzerkrankung. Derm. Wochenschr. 87: Wl-940. 1928. Ricketts, II. T. ('01). Oidiomycosis (blastomycosis) of the skin and its fungi. Jour. Med. Res. 6: 374-547. 1901. ■ — — , ('01a). A new mould fungus as the cause of three cases of blastomycosis or oidiomycosis of the skin. Boston Soc. Med. Sci., Jour. 5: 453-459. 1901. Roncali, I). B. ('95). Die Blastomyceten in den Sarkomen. Centralbl. f. Bakt. 18: 432-434. 1N<>5. Ryerson, E. W. ('08-'09). Blastomycosis: Report of two cases resembling bone tuberculosis. Am. Jour. Orthoped. Surg. 6: 79-83. 1908 1909. Sanderson, E. S., and D. C. Smith ('27). The effect of gentian-violet on the organ- ism of blastomycosis infection. Arch. Derm. & Syph. 16: 153-155. 1927. Sanfelice, F. ('95). Ueber einen neuen pathogenen Blastomyceten, welcher inner- halb der Gewebe unter Bildung kalkartig aussehender Massen degeneriert. Centralbl. f. Bakt. 18: 521-526. 1895. , ('96). Ueber die pathogene Wirkung der Blastomyceten. I. Abhand- lung. Zeitschr. f. Hyg. 21: 32 58. 1896. , ('96a). Ibid. II. Abhandlung. Ibid. 390-420. 1896. Schlossman, C. R. ('29). Two cases of blastomycosis cutis. Acta Dermato- Venereol. 10: 83-94. 1929. Simoni, A. de ('97). Ueber das Vorkommen von Blastomyceten in der Hyper- troplnschen Tonsille. Centralbl. f. Bakt. 22: 120-122. 1897. Smith, D. C, H. C. Turner, and E. S. Sanderson ('28). Systemic blastomycosis with a report of a fatal case. Brit. Jour. Derm. 40: 344-359. 1928. Speroni, I)., J. Llambias, S. E. Parodi, y. F. L. Nino ('29). Blastomicosis humano generalizado por criptococo (n. sp.). Efltudio parasitologic^), andtomopato- I6gica, elinico y experimental. Reuni6n Soc. Argentina Patol. Reg. Norte 5: 94-155. 1929. Spring, D. ('29). Comparison of seven strains of organisms causing blastomycosis in man. Jour. Inf. Dis. 44: 169-185. 1929. Stearn, E. W., and A. E. Stearn ('29). Comparative inhibiting effect of gentian violet and mercurochrome on the growth of certain fungi. Jour. Lab. & Clin. Med. 14: 1057-1060. 1929. Stober, A. M. (Ml). Systemic blastomycosis. Arch. Int. Med. 13: 509-556. 1914. Stovall, W. D., and II. P. Greeley ('28). Bronchomycosis. Report of eighteen cases of primary infection in the lung. Am. Med. Assoc, Jour. 91: 1346 1351. 1928. Sugden, F. ('23). Case of blastomycosis. Brit. Med. Jour. 2: 63. 1923. 1933] MOORE — BLASTOMYCOSIS 1 15 Sutejew, G., M. Utenkow, and A. Zeitlin ('29). Beitrag zur Atiologie, Rontgen- diagnose und Rontgentherapie der Blastomykose. Fortschr. Geb. Rontgenstr. 11:475-483. 1929. T., F. E. ('28). Cutaneous moniliases. Trop. Med. & Hyg. Jour. 31 : 37-38. 1928, Toepel, T. ('29). Systemic blastomycosis. Am. Med. Assoc, Jour. 93: 32. 1929. Tokishige, H. ('96). Ueber pathogene Blastomyceten. Centralbl. f. Bakt. 19: 105-113. 1896. Troisier, E., et P. Achalme ('93). Sur une angine parasitaire caus6e par une levure et cliniquement semblable au muguet. Arch. Med. Exp£r. 5: 29-37. 1893. Urbach, E., und F. Zach ('30). Generalisierte Torulose (Europaeische Blastomy- kose). Eine klinisch-botanisch Studie. Arch. f. Derm. u. Syphil. 162 : 401-421. 1930. Vuillemin, P. ('01). Les blastomycdtes pathog&ies. Rev. G6n. des Sci. 12: 732- 751. 1901. , ('10). Materiaux pour une classification rationelle des Fungi Imperfecti. Compt. Rend. Acad. Paris ISO: 882. 1910. Wade, H. W. ('16). A variation of gemmation of Blastomyces dermatitidis in the tissue lesion. Jour. Inf. Dis. 18: 618-629. 1916. ('18). Portal of entry in experimental chronic pulmonary (systemic) blastomycosis. Philipp. Jour. Sci. 13: 271. 1918. systemic , and G. S. Bell ( , 16). A critical consideration of Arch. Int. Med. 18: 103. 1916. Walker, J. W., and F. H. Montgomery ( ? 02). Further report of a previously re- ported case of blastomycosis of the skin: Systemic infection with blastomyces; death; autopsy. Am. Med. Assoc, Jour. 38: 867-871. 1902. Wanamaker, T. ('28). A case of blastomycosis of the cervical lymph gland. Am. Laryng., Rhin. & Otol. Soc, Trans. 34: 450-452. 1928. Weidman, F. D., and H. R. Douglas ('21). Blastomycetoid bodies in a sarcoma- like tumor of the leg. Arch, of Derm. & Syphil. 3: 743-752. 1921. Wernicke, R. ('92). t)ber einen Protozoenbefund bei Mycosis fungoides (?). Centralbl. f. Bakt. 12: 859-861. 1892. Whitman, R. C. ('13). A contribution to the botany of the organism of blastomy- cosis. Jour. Inf. Dis. 13: 85-94. 1913. Wilhelmj, C. M. ('25) . The primary meningeal form of systemic blastomycosis. Am. Jour. Med. Sci. 169: 712-721. 1925. Wohl, M. G. ('23). Fungous diseases of man in the State of Nebraska; sporotri- chosis; blastomycosis; actinomycosis. Am. Med. Assoc, Jour. 81: 647-653. 1923. Yakimoff, W. L., and W. J. Wassilewsky ('25). Au sujet de la blastomycose. Soc. Path. Exot., Bull. 18: 130-132. 1925. Zoon, J. J. ('30). Blastomycosis cutis durch Monilia floccoi mit positiver Blutkultur. Derm. Wochenschr. 58: 35&-367. 1930. [Vol. 20, 1933] 116 ANNALS OF THE MISSOURI BOTANICAL GARDEN Explanation of Plate PLATE 6 All drawing! made with camera lucida at a magnification of X 800. Figs. 1-9. Yeast-like cells. Figs. 1, 6, 7. On Raulin's solution. Figs. 2-5, 9. On Sabouraud's agar. Fig. 8. On glycerine agar. Figs. 10-12, 14-17. Yeast-like cells showing a change to mycelial formation. Figs. 10, 12. On potato-dextrose agar. Figs. 11, 14. On glycerine agar. Figs. 15-17. On Sabouraud's agar. Fig. 13. Germinating spores on Richards' solution agar. Fig. 18. Heterogarnous copulation of lateral cells on Sabouraud's agar. Fig. 19. Heterogarnous copulation of terminal cells on Richards' solution agar. Fig. 20. Copulating branch on corn-meal agar. Fig. 21. Maturing ascus on Richards' solution agar. Fig. 22. Terminal hypnospore on Sabouraud's agar. Fig. 23. Racquet mycelium on June-beetle agar. Fig. 24. Mycelium showing conidia on Richards' solution agar. Fig. 25. Mycelium showing round terminal chlamydospores and swollen hypha on nutrient agar. Figs. 26, 28. Mycelium showing conidia, oidia-like cells, and resting cells on corn-meal agar. Fig. 27. Chlamydospore on Czapek's agar. Fig. 29. Mycelium showing conidia on potato-dextrose agar. Fig. 30. Racquet formation on Czapek's agar. Fig. 31. Terminal chlamydospore on lactose agar. Fig. 32. Terminal hypnospore on Endo's agar. Fig. 33. Maturing lateral ascus on Sabouraud's agar. Fig. 34. Racquet mycelium on June-beetle dextrose agar. Fig. 35. Ascus covered with a third sheath in proximity to a round resting cell, on potato-dextrose agar. Fig. 36. Lateral chlamydospore on Czapek's agar. Fig. 37. Resting cell on Sabouraud's agar. Fig. 38. Mature ascus on potato-dextrose agar. Ann. Mo. But. Gard., Vol. 20, 1933 PliATE 6 MOORE— BLASTOMYCOSIS [Vol. 20, 1933] 118 ANNALS OF THE MISSOURI BOTANICAL GARDEN Explanation of Plate PLATE 7 Fig. 1. Photograph of hand of patient on day of entry, April 8, 1932, showing lesion involving portion of thumb. Fig. 2. Photograph taken on April 8, 1932, showing abscess on flexor surface of left lower arm. Fig. 3. Photograph showing marked improvement after treatment with sodium iodide intravenously. Fig. 4. Photograph showing almost complete healing. Ann. Mo. Bot. Gaud., Vol. 20, 1933 Plate 7 fZbU 050095 MOOKE— BLASTOM YCOSIS A MONOGRAPH OF THE AMERICAN SPECIES OF THE GENUS HALENIA 1 CAROLINE K. ALLEN Formerly Missouri Botanical Garden Special Fellow in Botany^ Henry Shaw School of Botany of Washington University Introduction Frequent attempts to determine recent collections of Halenia, especially from Central and South America, have revealed the need for a comprehensive taxonomic treatment of the American species of the genus. Incidental determination of isolated species in herbaria, which furnish scanty material, has led to error and the duplication of species has resulted. Few of these fragmentary treatments are provided with adequate descriptions, and still fewer are accompanied by illustrations. The present paper is a monographic study of the American species of Halenia. The first portion is devoted to the North and Central American representatives of the genus and the second to those of South America. The writer at this time wishes to express her appreciation to Dr. George T. Moore, Director of the Missouri Botanical Garden, where this work has been carried on, for the privilege of using the facilities afforded by the herbarium, library, and laboratory. It is with pleasure, indeed, that she also acknowledges the courte- ous assistance and helpful criticism offered by Dr. J. M. Green- man, under whose personal supervision the study was begun and completed. For the loan of specimens for study the writer is especially indebted to the curators of the herbaria of the following institu- tions: Gray Herbarium of Harvard University, New York Botanical Garden, Philadelphia Academy of Natural Sciences, United States National Herbarium, Field Museum of Natural History, the Dudley Herbarium of Stanford University, Brooklyn Botanic Garden, Iowa Agricultural College, Natural History Mu- 1 An investigation carried out at the Missouri Botanical Garden in the Graduate Laboratory of the Henry Shaw School of Botany of Washington University, and submitted as a thesis in partial fulfillment of the requirements for the degree of doctor of philosophy in the Henry Shaw School of Botany of Washington University. Issued April 29, 1933. Ann. Mo. Bot. Gard., Vol. 20, 1933. (119) 120 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 20 seum of Vienna, Botanical Museum of Stockholm, the Delessert Herbarium at Geneva, Royal Botanic Gardens, Kew, and the British Museum of Natural History. Particularly does the writer wish to express her gratitude to Mr. R. I. Cratty, of the Iowa State College, for the loan of specimens from the Parry Herbarium deposited there; to the curator of the Herbarium of the Botanical Garden of Madrid, for supplying a photograph of the type of Swertia cucullata; and to the curator of the Herbarium of the Jardin des Plantes, Paris, for photographs of the types of Humboldt, Bonpland and Kunth. In connection with visits to various herbaria, the author desires to acknowledge the courtesy and kindly assistance of Dr. Ernst Gilg and Dr. Robert Pilger, of the Botanical Garden at Berlin-Dahlem ; Dr. Walter Robyns of the Botanical Garden, Brussels; and Mr. Spencer Savage, in charge of the Linnaean Herbarium, of the Linnaean Society of London. Thanks are due also to Dr. George E. Nichols, Director of the Marsh Botanical Garden, Yale University, for his kindness in procuring seeds of Halenia; to Miss Nell C. Horner, Librarian of the Missouri Botanical Garden, and Dr. John H. Barnhart, of the New York Botanical Garden, for assistance in bibliography; to Dr. Roland V. La Garde, of the Missouri Botanical Garden, for preparation of photographs; and to the George F. Cram Com- pany, for permission to use the copyright outline maps of North and South America. History of the Genus Linnaeus in the 'Amoenitates Academicae/ 1 which appeared in 1751, published short descriptions of two genera of the Genti- anaceae, namely, Swertia, consisting of five species, and Gentiana, of twenty-three. Under Swertia Linnaeus listed the spurred gentian with the following description. "4. Swertia corollis quadrifidis quadricornibus. Amoen. acad. 2. p. 344. Habitat in Sibiria, Gmelin; Canada, Kalm." This was apparently the only species of spurred gentian he had 1 Linnaeus, Amoen. Acad. 2: 344. 1751. 1933] ALLEN — THE GENUS HALENIA 121 ever seen, and, it being closely related to the Swertia he knew, he placed it in that group, giving it the specific name corniculata or "horn-tipped," as distinct from the others. Gmelin, in his 'Flora Sibirica,' 2 published in 1769, referred to this as being synonymous with his genus Tetragonanthus, which he had described or mentioned in a previous book or manuscript, and which he evidently based upon specimens collected by G. H. Stellar. This work contains a brief description, but a good illustration of Swertia. On account of the presence of spurs on the corolla, Borck- hausen, 3 in 1796, segregated Swertia corniculata from the Linnaean genus Swertia and called it Halenia after Jonas Halen. Although the latter had included a short description of it in a previously published dissertation on Kamtchatka plants, Borckhausen must be considered the author of the genus. He cited as a synonym Swertia corniculata Linnaeus, but changed the binomial name to Halenia sibirica. The name Swertia, however, persisted for some time in litera- ture, the generic descriptions becoming more elaborate and detailed with each publication. Ruiz and Pa von described and illustrated Swertia umbellata from Peru 4 in 1802. Michaux included the genus in his 'Flora Boreali- Americana ,fi appearing the following year, and Humboldt, Bonpland and Kunth, 8 described six new species from Mexico and South America in 1818. A few species, together with a new variety, were published in 'Linnaea' by Schlectendal and Chamisso 7 in 1830. But it was not until Grisebach began his extensive study of the Gentian- aceae that any attempt was made to bring together the species of the world. As a result, when his ' Observationes' 8 appeared in 1836, followed in 1839 by the 'Genera et Species Gentian- earum,' 9 many of the existing names fell to synonymy. Grisebach divided Halenia into two main divisions: the first, * Gmelin, Fl. Sib. 4: 114. 1769. 3 Borckhausen in Roemer, Arkiv fur Botanik l 1 : 25. 1796. 1 Ruiz & Pavon, Fl. Peru v. 3: 21. pi. 242. 1802. 6 Michaux, Fl. Bor.-Am. 1 : 97. 1803. 6 Humboldt, Bonpland & Kunth, Nov. Gen. & Sp. PI. 3: 174. 1818. 7 Schlectendal & Chamisso in Linnaea 5: 122. 1830. 8 Grisebach, Obs. Gent. 36. 1836. • Grisebach, Gen. & Sp. Gent. 322-328. 1839. [Vol. 20 122 ANNALS OF THE MISSOURI BOTANICAL GARDEN with spurs ascending and spreading; the second, with spurs pendulous to incurved. He devoted careful attention to the descriptions, synonymy, affinities, differentiating characters, and habitat of each species mentioned. The collector and type were given in each case. He also separated from the Swertia of Humboldt, Bonpland and Kunth two species, brevicomis and parvijfora, and founded on them a new genus Exadenus distin- guished by the presence of pits at the base of the corolla, instead of spurs, and by central placentation. This abolished the Swertia of Humboldt, Bonpland, and Kunth, which was based on Swertia corniculata Linnaeus, and left the non-spurred Swertia originally described by Linnaeus a genus entirely distinct from our present Halenia. Hooker's 'Flora,' 10 published in 1840, contained good illustra- tions of Halenia dejlexa with the varieties Brentoniana and heterantha. Nearly a decade elapsed before any significant study was done on the genus as a whole. Bentham, 11 in 1839-1840, in describing Hartweg's plants from Mexico added two new species, multiflora and decumbens. Martens and Galeotti in 1844 in their " Enu- meration of Mexican plants collected by Galeotti" 12 described, along with other new Halenia species, two new species of Exa- denus. Walpers' ' Repertorium ' 13 contains reference to Exadenus, but Weddell in 1859 14 merged the two genera. He considered the group as a whole to consist of two main subdivisions, the first being that group with spurs, and the second the spurless species. The former he subdivided into section 1 — ovary uni- locular; section 2 — ovary bilocular, and the corolla having small spurs. Here he placed the two original species of Exadenus Grisebach which were based on the species of Swertia Humboldt, Bonpland & Kunth, namely, yarviflora and brevicomis, but neglected to transfer the species described by Martens and Galeotti. The second subdivision, without spurs, he also divided into two sections, on the presence of a uni- or bilocular ovary. 10 Hooker, FL Bor.-Am. 2: 67. pi. 156-6. 1840. 11 Bentham, PI. Hartw. 24. 1839; 67. 1840. 12 Martens & Galeotti in Bull. Acad. Brux. 11': 370. 1844. 18 Walpers, Rep. Bot. Syst. 6: 508. 1840-47. m Weddell, Chlor. And. 2: 74. 1859. 1933] ALLEN — THE GENUS HALENIA 123 His argument for combining the two genera was based on the fact that the two characters, the bilocular ovary and the short spurs do not always coincide. In this work, he described five new South American species based on collections of Triana, Funck & Schlim, Goudot, Purdie, etc. At the time of the publication of Bentham and Hooker's 'Genera Plantarum,' 15 1876, there were about twenty-five rec- ognized species of Halenia from the whole world. Hemsley, in 'Biologia Centrali- Americana,' 16 which appeared in 1882, referred the Exadenus Martens and Galeotti to Halenia. Gilg in Engler and Prantl's 'Naturlichen Pflanzenfamilien' 17 recognized the validity of the generic name Halenia and included a careful description and good illustrations. Sess6 & Mocino's 'Flora' 18 contains a description of Swertia cucullata which has escaped the notice of succeeding monog- raphers, and has not been transferred to Halenia. A photo- graph of the type specimen kindly furnished by Dr. E. Balguerias, Curator of the Botanical Garden at Madrid, reveals the fact that it is without question Halenia brevicornis Griseb., though it is rather difficult to determine whether or not it is the species proper or one of its many forms. From this time until the monographic treatment of South American species was published by Gilg 19 in 1916, several species and varieties of Mexican and Central American Halenia were published. Halenia Rothrockii Gray, 20 now Halenia recurva, in 1876 was based on plants collected in Arizona by Dr. J. T. Rothrock. G. Don, in 1838, in the 'General History of the Dichlamydeous Plants' 21 included a description of the existing species of Halenia, making several new combinations from the species of Humboldt, Bonpland and Kunth. 22 Some of his species were taken from the manuscript of D. Don, thus appearing in 16 Bentham & Hooker, Gen. PL 2: 817. 1876. 16 Hemsley, Biol. Cent.-Am. Bot. 2: 351. 1882. 17 Gilg in Engler & Prantl, Nat. Pflanzenfam. 4 2 : 89. 1895. 18 Sesse & Mocino, Flora Mexicana, 73. 1894. 19 Gilg in Engl. Bot. Jahrb. 54: Beibl. 118, pp. 93-122. 1916. 20 Gray in Proc. Am. Acad. 11 : 84. 1876. » Don, G. Gen. Hist. 4: 177. 1838. 22 Humboldt, Bonpland & Kunth, Nov. Gen. & Sp. PI. 3: 174. 1818. 124 ANNALS OF THE MISSOURI BOTANICAL GARDEN (Vol. 20 publication for the first time. Kuntze, 23 in 1891, revived the generic name Telragonanthus Gmelin. He has been followed by a few later taxonomists, among whom are Britton, 24 Small, 26 Rydberg, 26 and others. However, Halenia was placed on the list of nomina conservanda , and thus must be regarded as the correct name in accordance with the International Rules of Botanical Nomenclature. Halenia crassiuscula Robinson and Seaton 27 ap- peared in 1893. Britton, 28 in 1894, reduced H. Brentoniana Grisebach to a variety of Tetragonanthus deflexus, while in 1899 Fernald 29 placed H. heterantha Grisebach under Halenia deflexa var. heterantha. Other new species and new combinations ap- peared in rapid succession, due to the extensive collecting done during the period from 1890 to about 1920, by Conzatti, Pringle, Purpus, and others in Mexico, and Weberbauer in South America. Among these newly published species were: Halenia Candida Ramirez, 30 1895; H. chlorantha Greenman, 31 1905; H. bella, H. caespitosa Gilg, 32 1906; H. Conzattii Greenman, 33 1912; H. guate- malensis Loesener, and H. plantaginea var. latifolia Loesener 34 (now //. guatemalensis var. latifolia (Loesener) Allen) 1913. The type species up to this time had been called H. sibirica, the name given it by Borckhausen. However, according to the rules of priority, sibirica should give way to the older name corniculata used by Linnaeus. Accordingly, Druce 38 in 1914 revived corniculata which is at present the accepted name for the type species of Halenia. Britton and Brown in the 'Illus- trated Flora,' 36 reduced Grisebach's species Brentoniana and heterantha to varieties of deflexa. M Kuntze, Rev. Gen. PI. 2: 431. 1891. u Britton, Manual, 734. 1901. « Small, Fl. Southeastern U. S. 931. 1913. M Rydberg, Fl. Rocky Mts. 666. 1922. 17 Robinson & Seaton in Proc. Am. Acad. 28: 113. 1893. " Britton in Mem. Torrey Bot. Club 5: 261. 1894. " Fernald in Rhodora 1 : 37. 1899. 10 Ramirez in Inform. Secret. Foment. Mexic. (Excurs. Mont. Ajusco). 34. 1895. B Greenman in Proc. Am. Acad. 41: 240. 1905. « GUk in Fedde, Rep. Spec. Nov. 2: 52. 1906. » Greenman in Publ. Field Mus. Bot. 2: 335. 1912. u Loesener in Verh. Hot. Ver. Brandenb. 55: 182. 1913. * Druce in Rept. Bot. Exch. CI. Brit. Isles 3: 419. 1914. 16 Britton & Brown, 111. Fl. 3: 15. 1913. 1933] ALLEN — THE GENUS HALENIA 125 Gilg, 37 in 1916, monographed the South American representa- tives of the genus, adding several new species. At that time, the herbarium at Berlin contained as complete a series of South American plants as were to be found anywhere, but, in some cases at least, the material was too inadequate to determine the limits of variation of certain of the species proposed. Hence, some of the species recognized, studied in the light of subsequent collec- tions, have fallen into synonymy. Gilg created three main divisions : A. Nectaria parva vel obsoleta, rarius extrinsecus breviter semigloboso-prominentia. Folia manifeste carnoso-sub- coriacea. B. Nectaria haud calcariformia, sed extrinsecus ad basin corollae alte globoso- vel coniformi-prominentia. Folia semper tenuiter herbacea. C. Nectaria extrinsecus calcaria manifeste evoluta formantia. Division A contains the largest known Halenia, native of South America only, which in inflorescence, flower, and leaf habit, shows affinities with the western species of Swertia and Frasera as well. Plants in division A are very distinct and occur in Colombia and Venezuela. These Gilg apparently considered the most primitive. In division B we find the brevicornis-parviflora complex. Gilg considered H. brevicornis (HBK.) Don a valid species known only from South America, and parviflora a native of Mexico; he fur- ther described a new species erythraeoides from Venezuela which agrees in every respect with the parviflora type from Mexico. Division C is subdivided according to the length of spurs ; here again, confusion has occurred, as is apt to be the case when herbarium material is scanty and field work impossible. From a limited experience in the field with a North American species, the author can state with conviction that it is possible for two plants belonging to the same species, growing side by side, to vary not only in form but in size as well. It has been found also that the spur character is inconstant. Axillary flowers and those blooming late in the season frequently possess shorter spurs or no spurs at all. If this character is so variable in one species, it 17 Gilg in Engl. Bot Jahrb. 54: Beibl. 118, p. 93. 1916. 120 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 20 is reasonable to suppose that the same situation may obtain in other species. Only close attention to ecological detail and a wealth of material not yet available from South America can enable the monographer to delimit species from these little- known regions even with a fair degree of accuracy. Briquet, 38 in 1931, described several new species which will be treated below. Gross Morphology Habit. — The American species of the genus Halenia are gla- brous, strictly herbaceous, or somewhat ligneous annuals or per- ennials. They may be of caespitose habit, as is illustrated by H. caespitosa, or coarse, fleshy, foliose plants with single stems, 5 or more dm. high, as in H. hygrophila and related species, or scapose with 1 lie basal leaves disposed in a rosette, as in H. plantaginea , or slender, graceful, simple or branching plants 1.5-6 dm. high, as in //. brevicornis and its varieties. As a whole, the genus is not colorful, the flowers being various shades of yellow and yellow- green, except in the northernmost species, which has purple flowers. Roots. — The root system of the North American species is fibrous, with a persistent, slender tap-root, frequently more or less woody in texture, though typically that of an annual or perennial in cross-section. In the South American species the root is, for the most part, ligneous and thick. Stems. — The stems are simple or branched, usually erect, though they may be decumbent, as in Halenia decumbens and H. Weddtlliana. They are of two types, mostly foliose as in H. deflexa and Schiedeana, or scapose as in H. plantaginea. The stems often continue underground for a short distance, sending out erect, flowering stems at irregular intervals. These may be angular or terete; if the former they are often slightly winged, due to the decurrence of the leaves, and usually faintly striate. Leaves. — The leaves are opposite and decussate, or infrequently whorled as in H. verticillata , either entirely cauline or disposed in a rosette, sessile or petiolate. The petioles exhibit the same characteristics as the stem, the decurrence of the calyx-lobes M Briquet in Candollea 4: 317. 1931. 1933] ALLEN — THE GENUS HALENIA 127 frequently being apparent. Where both petiolate and sessile leaves occur on the same plant, the basal leaves have petioles often equal to or longer than the blade, and the petioles become decreasingly shorter toward the summit of the stem, the upper- most leaves being sessile. The leaves are entire, 1-, 3- or 5- nerved, and range from ovate to obovate or spatulate, or lanceolate to linear, varying from .5 to 8 or 10 cm. in length and up to 6 cm. in width. The leaves of the North American species are, on the whole, thin and herbaceous, whereas in the South American species they are frequently coarse, fleshy, or coriaceous. Inflorescence. — The inflorescence consists of terminal or axillary cymes of varying density. The flowers occur on short pedicels at the tip of a stem or branch in a compact cluster or head, as in H. brevicornis, or they may be borne loosely on long pedicels at the tip of a branch or at a node, as in H. Schiedeana. In the Central American and the more primitive of the South American species, the inflorescence is usually a spicate, racemose, or more or less umbellate cyme. A pair of small, linear, foliaceous bracts is usually present at the base of the inflorescence, often bearing tiny, undeveloped buds in their axils. Frequently the bracts approximate the leaves in size and structure, and in H. involucrata they form an involucre almost entirely enveloping the inflores- cence. Calyx. — The calyx is persistent, foliaceous, 4-parted, and the segments united only at the extreme base and arranged in pairs; the inner pair represents an inner cycle, the second an outer cycle. The segments may be caudate as in the type species, H. corni- culata, or spatulate, obovate, elliptic, lanceolate-linear, or linear, with intergradations, with obtuse, acute, acuminate, mucronate or apiculate tips. The tips very rarely are reflexed, as in H. Schiedeana. The length of the calyx varies from one-third to that egments are from 1- to 3-nerved, often reticulately veined at the tip, and both surfaces are often papillate. Squamellae are usually found on the inner surface of the calyx-segments at their base (see fig. 1). They vary in size, shape, number, and position on the lobes, and 12S [Vol. 20 ANNALS OF THE MISSOURI BOTANICAL GARDEN are usually distinguishable only under the dissecting lens after the herbarium specimens have been boiled. Squamellae occur in many of the Contorti both on the calyx and the corolla-lobes. Engler and Prantl 39 have referred to similar structures in several genera of the Gentianaceae as discs. Fig. 1. Types of squamellae found on the calyx in the genus Halenia. Corolla. — The corolla is marcescent, campanulate, 4-lobed, white, yellow, yellow-green, green, or purple, ranging in length from 5 mm. to nearly 3 cm. The lobes are triangularly ovate or obovate, acute, acuminate, apiculate, mucronate or obtuse, often auriculate, with an entire, erose, or crisped margin, and fre- quently papillate on both surfaces. The veining of the dex- trorsely convolute lobes may or may not be reticulate. The corolla-tube varies from one-fourth to three-fourths the length of the entire corolla. At or near the base of the tube, opposite each lobe, is a tubercle which may be merely a slight swelling in the lower portion of the tube, as in H. brevicornis, or a definite 3 » Gilg in Engler & Prantl, Nat. Pflanzenfam. 4 2 : 89. 1895. 1933] ALLEN — THE GENUS HALENIA 129 spur longer than the corolla, as in H. guatemalensis (pis. 8-11). It frequently happens that the corolla lacks spurs entirely in the axillary flowers or in those occurring late in the season, as, for example, in H. defiexa and other species. Not only in Halenia is this situation apparent, but it is recalled that in other normally spurred plants, for example, Linaria canadensis, spurs are fre- quently absent. Therefore it has seemed advisable to discon- tinue heterantha as a variety of H. defiexa, since the occurrence of these spurless forms is more or less frequent within the genus. Gilg noted that in certain species of Halenia, for example, H. brevicornis, etc., the flowers on the main stalk are normally large and wide open, while below, on the same stem, they are definitely smaller, apparently not opening at all or else very slightly. The structure is similar, with the exception of the absence of spurs. The ovary in these abnormal flowers contains fewer seeds, and the capsule is much reduced in size. This would indicate, according to Gilg, a gradual reduction of chasmogamous flowers which depend on insect pollination, to more or less cleistogamous flowers. The shape of the spurs varies exceedingly, from slender to very thick and broad, or from spreading and ascending to pendulous and incurved. Intergrading forms are frequent. The spurs are frequently conspicuously veined and apparently glandular. Stamens. — The stamens are equal in number to, and are borne alternate with, the lobes of the corolla at or near the summit of the tube. The filaments are adnate to the corolla, and the anthers are often enfolded in the bud by the margins of the corolla-lobe. The anthers are versatile, deltoid, ovate or oblong, often mucronate and papillate. The filaments are usually linear, but they may become more or less dilated and, rarely, papillate. The pollen grains are constant for the genus, being more or less tetrahedrally spherical, and having three pores. Pistil. — The pistil is sessile. The stigma is usually sessile and cleft to expose the two inner stigmatic surfaces ; the lobes may be truncate or ovate. The ovary is bicarpellate, the margins of each carpel being infolded and becoming the region of attachment for the numerous ovules. Fruit. — The fruit is a flattened, lanceolate to lanceolate- 130 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 20 obovate capsule, frequently subfalcate, and usually exserted. It is unilocular at maturity, though in the young state it appears to be more or less two-celled. The fruit dehisces septicidally along the inner surface of each locule tip. Seed. — The seeds vary in shape, being globose to ovoid or elliptic, often flattened. The surface is reticulate in the majority of the South American species and in the brevicornis complex, except for var. latifolia. The remaining species show the surface of the seeds to be minutely granular instead of reticulate. They vaiy in size from 0.5 to 1 mm. in diameter, and in color from dull greenish-brown and yellow-brown to dark, shiny brown, the latter usually being typical of those with reticulate surface. The age of the plant and the conditions attending its collection no doubt have their influence on the color, size, and, to some extent, the texture of the seed-coat. Floral Anatomy The major portion of the anatomical investigation of the Gentianaceae, particularly the Menyanthoideae, has been con- cerned with the stem and leaf structure. The most complete anatomical data is found in Gilg's treatment of the Gentianaceae in 'Die Naturlichen Pflanzenfamilien.' 40 Solereder's 'Systematic Anatomy of the Dicotyledons' 4 ' gives very little additional in- formation. Since that time more attention has been given to floral morphology as a separate study. Stolt, 42 in 1921, made an exhaustive cytological survey of the flowers of several genera, among which was included Halenia elliplica, an Asiatic species having affinities with Halenia dejlexa of North America. Inci- dentally, Stolt inserted a diagram of the transverse section through the ovary, which indicates clearly the vascular system of that portion of the flower. In order to make the present monograph as complete as possible, anatomical study was undertaken. Fresh flowers of Halenia deflexa, a species with both spurred and spurless forms, were obtained by the author in Vermont and preserved in 70 per cent <° Gil* in Engler & Prantl, Nat. Pflanzenfam. 4 2 : 50. 1895. 41 Solereder, Systematic Anatomy of the Dicotyledons 1: 548-550. 11)08. "Stolt in K. Svensk. Vet.-Akad. Handl. 61'*: 1-56. 1921. 1933] ALLEN THE GENUS HALENIA 131 alcohol. The pickled material was dehydrated and embedded in paraffin following the butyl alcohol method outlined by Zirkle, 43 sectioned at 10 (i, and stained with crystal violet and erythrosin. The accompanying drawings (pi. 12) were made with the aid of a "Promi" microscopic drawing and projecting appara- tus. The xylem has been cross-hatched in order to differentiate it from the other vascular elements. Transverse sections of the spurred form of Halenia deflexa show that the vascular system of the pedicel is an amphiphloic siphonostele (pi. 12, fig. 1). Approaching the receptacle, the stele enlarges and assumes a rhombic form (fig. 2). The de- currence of the outer lobes of the calyx is apparent. Shortly thereafter (fig. 3), the midribs (a) of the two outer calyx-lobes leave the receptacular stele (r). They migrate outward, and from either, two lateral traces (a') are given off (fig. 4) . At this point, four lacunae (z) appear, prior to the severing of the calyx from the receptacle. In the succeeding illustration (fig. 5) these lacunae (z) merge into two crescent-shaped fissures, and the midribs of the two inner calycine lobes (b) leave the stele. In fig. 6 the calyx-tube is entirely free from the receptacle, and in the axil of each potential lobe are visible 4-8 minute emergences or squamellae (e). The stele has again assumed a more or less rhombic shape, but from the 4 angles, traces (c) depart centri- fugally, soon resolving into the midrib (n) and two laterals (c') destined to supply each of the 4 corolla-lobes (fig. 7). The disruption of the residual vascular cylinder is continued, and the 4 staminal traces (f) are fully differentiated. At this in- terval the corolla-tube is virtually free from the receptacle. In the succeeding illustration (fig. 8) lacunae appear at m, the ovary at this point appearing bilocular. The residual stele con- sists of two roughly semi-circular masses with a concentration of lignified elements (1) at either end. The calyx-lobes are free, and sections near the tip of the reflexed spurs are found, the origin of which is to be described. In the following illustration (fig. 9) the origin of the spurs is apparent, and a cleft (y) is visible, indicating the sinus. Simul- taneously, 4 protrusions occur on the inner surface of the corolla, « Zirkle in Science, N. S. 71 : 103. 1930. 132 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 20 below the sinus, preparatory to the severing of the staminal filaments (f). The ventral traces (1) and dorsal (k) are dis- cernible. The ovules (o) are evident, showing their position in the axils of the placentae (p). In the final stage, taken from a section through the tip of the flower (fig. 10), the differentiation of the corolla-lobes and the isolation of the staminal filaments (f) are complete. The pla- centae (p) have diminished in size. The spurless form shows a similar vascular system except for the absence of spurs, the presence of fewer ovules, and finally, a more pronounced dorsal trace (k) , after the cessation of ovular production. Geographical Distribution The accompanying maps show three centers of distribution of the species of Halenia in America. The first (figs. 2, 4) extends from Labrador and Newfoundland, south to New York and west to British Columbia and Montana. The one species and its variety found in this area grow in moist or dry situations in calcareous, slaty, or alluvial soil, in open woods or fields, on stream banks or along the sea-shore, usually in the shade. The habit varies with the habitat. This distribution follows closely the northern region of glaciation, and coincides with the usual distribution areas of herbaceous species common in that territory. This same species has been collected three times in the State of Mexico, but has not been reported from the intervening region, a fact that might suggest a previously more continuous distribu- tion of the species from the northern Rockies along the mountain ranges to the Mexican Sierras. The second area comprises a region extending from the Chiri- cahua Mountains of New Mexico and Arizona, southward to Costa Rica in Central America. Here are a few wide-spread mountain species; but for the most part, they are endemics occurring in volcanic areas (fig. 3). The third large center of distribution is the northern part of South America, where the genus is represented by a relatively large number of endemic species (fig. 5). All of the North and Central American species, with the Fig. 2. Halenia H. deflexa; 2 = H. deflexa var. Brenioniana. 14. H. brevicornis Fig. 3. Map of southern United States, Mexico, and Central America showing the geographical distribution of Halenia. 19. 18. 15. 17. var. chihuahvensis var. divergent var. latifolia var. micranthdla 16. 20. 21. var. mvltiflora var. ovata var. Tuerckheimii 29. //. caleoides 6. H. Conzattii 4. H. crassiuscida 13. H. decumbens 25. ff. guatemalensis 30. var. latifolia 12. //. nudicavlis 5. #. Palmeri 8. i/. plantaginea 9. f. grandiflora 22. tf. platyphyUa 10. #. Pnnffta 3. //. recurva 26. //. rhyacophila 28. 27. 24. //. Shannonii 23. f. compncta var. procumbens var. macro pod a 11. //. Schiedeana 1933] ALLEN THE GENUS HALENIA 133 6 7d p O o CO •2 I P O P C .2 i3 a '3 •B - O go a* to o P o CO fl .P GO O t« I CO J3 exception of Halenia alata and iL brevicornis and its varieties, are spurred forms which show a definite relationship to one another. Those from South America are of two types: spurred type, similar to the North American species (excluding brevicornis and 134 ANNALS OF THE MISSOUKI BOTANICAL GARDEN [Vol. 20 alata) and showing affinities with them; non-spurred type, with tubercles instead of spurs and quite different in habit from the brevicornis and alata. Halenia brevicornis and its varieties form a connecting link between the second and third areas, as well as a morphological link, so to speak, between the primitive Swertiella and the more advanced Haleniastrum. The varieties of brevi- cornis do not extend further south than Guatemala, though the species proper is found in South America. Here again, as in Mexico and Central America, exist endemics which show a development almost parallel with that which has occurred north of the equator. An example of this is clearly shown by Halenia decumbcns, from the mountains of Mexico, which bears a striking resemblance to //. Weddelliana. a species from Ecuador, Colombia, d Per Systematic Position The Gentianaceae, as pointed out by Gilg, 44 shows closer affinity to the Loganiaceae than to any other family in the Contorti, yet the differences are so well marked that the two families are never confused. Grisebach separates the Gentianaceae into two subfamilies, the Gentianoideae and the Menyanthoideae. In the first subfamily the leaves are always simple, entire, sessile, and never al- ternate. The aestivation of the corolla is never valvate. The second subfamily, the Menyanthoideae, has alternate and mostly petiolate, sometimes trifoliate, leaves, and the aestivation is induplicate-valvate. Halenia belongs to the first division. The structures called squamellae, described early in the text, are never found in the Menyanthoideae but may occur in other genera of the Gentianaceae, at the base of the corolla or calyx- lobes. The nearest relative of Halenia is Swertia. The most primitive forms of Halenia, particularly the South American species, are often confused with Swertia. The primitive members of Halenia, instead of the definitely spurred corolla typical of the majority of the species of the genus, possess small, knob-like protuberances or nectaries which upon casual examination might pass unnoticed 44 Gilg in Engler & Prantl, Nat. I'flanzenfam. 4 2 : 60. 1895. 1933) ALLEN — THE GENUS HALENIA 135 or be taken for the nectaries which are structurally distinct and characteristic of Swertia. More detailed study reveals the fact that the depressions or spurs of Halenia are without the marginal fringe which is always conspicuous in Swertia (fig. 6). Halenia present. The e in the bud. has a corona, but in Swertia it is frequently lobes of Halenia dextrorsely whereas the reverse is the case in Swertia discloses other generic differences. Anatomical study a c Fig. 6. Swertia perennis L. : a, bud; b, interior surface of petal. (HBK.) Don: c, bud; d, interior surface of petal. brevicornis Within the genus itself two rather distinct parent. The more primi which was m abov as Swer being near Swertia and possibly originating from it, may be called tentatively Swertiella. It contains those species from South America which were considered by Gilg as being most primitive. Halenia alata of Mexico also belongs to this section. The tiellae may be characterized by the absence of spurs or the pres- ence of small tubercles or prominences at the base of the corolla, which are not visible usually from the outside. They are coarse, fleshy, foliose plants, and for the most part with dense inflores- cences. The second section, H aleniastrum , is more advanced and is readily recognized by the presence of spurs of varying (Vol. 20 136 ANNALS OF THE MISSOURI BOTANICAL GARDEN length and shape. Both herbaceous and more or less woody types comprise this section. An interesting discussion of the phylogeny of the Gentianaceae was presented by T. H. Huxley 45 before the Linnaean Society of London, April 7, 1887. Confining his study to the structure of the corolla, he separated the family into two groups; the first division Permelitae was segregated mainly on the presence of a series of inner surface of The Permelitae were again divided into four groups with various PUITTKVliA 0UA]U (A1£,Ci > IS KYTralCOIDIS mo: PUNT AS niKA rni i roiOAi PREVICORIUS KARSTKK1 OTHLSU* BACIIAfC.HA If* lAHOIIfc STTTULHIOUBS V1RIDIS— 1 T0LIMAI I SUB IK VO LUC TTUl^UALIS 1 ' SCHULTZEI ' MAJOR VEHTTCrLlATA HTOROPHrLA JRASUIA oWXRTIA _1 PARALIZLA DASYANTPA KLATa 'ONI I PURDtEAffA RUSBYI H Hisnoima BARB ICADXI & PRBRUAI PA1MLRI ASCtXI UlffiA' RHYACOPH HA1 I ZX5GA f~~ HOPPI RBCURVA p, ZISGANS I CONZATTII OXDES — DECUlSiHS L-WKDiailXlAHA' |-BI*IHA KAXBRIYERI BELLA SPHAQSICOLA • «•! CIXIS VALKRlANOrrES -CRASS IUSC ULA-CAESPITOSA PUS ELLA SPATULATA HERZOGII SLLi?*)IDJ8 -IGANTSA MINIMA WEBJvRBAUER I pKYTsir-roiras-j I psiroirLinflRA MJYLRI JOHAM*IS«l I " MSi *£LA T A I: IKCETOXICffrOES ROEUSTA MATHER SI I STUJBJXII I • ga (•/**_> KYTOTKET IC At AMCKTPR Fig. 7. Chart showing the probable phylogenotic relationship of species of Halenia modifications of Actinanthe, which he lered the most primitive and the least differentiated tained among other elements, Exade eluded as a transitional stage on accoun leading to the second division, the The latter was in- vision, 11h> Kvratanihe, containing only Halenia. which represents an extreme modification of Exadenus The third related division, the « Huxley, T. H. Jour. Linn. Soc. Bot. 24: 101. 1887. 1933] ALLEN — THE GENUS HALENIA 137 depicts another outgrowth of the Actinanthe type, but with filamentous appendages or fimbriae, Swertia and Frasera fall into this category. Text-figure 7 shows the probable relationship of the various species of Halenia, according to the author's interpretation. For the sake of convenience, in the taxonomic treatment of the genus, the North and South American species are taken up separately in each section. Abbreviations In the citations of specimens examined the following abbrevia- tions have been used to denote the various herbaria from which specimens were used for study. ANSP = Herbarium of the Academy of Natural Sciences of Philadelphia. BG = Botanical Garden, Berlin. BB = Brooklyn Botanical Garden. BM = British Museum of Natural History, London. B = Herbarium of the Botanical Garden, Brussels. C = Herbarium of the University of Chicago, deposited in the Field Museum of Natural History. CAS = Herbarium of the California Academy of Sciences. D = Dudley Herbarium of Leland Stanford Jr. University. DH = Delessert Herbarium of Geneva. F = Herbarium of the Field Museum of Natural History. G = Gray Herbarium of Harvard University. HP = Herbarium of H. Pittier, Director de Museo Comercial, Caracas, Venezuela. HJP = Herbarium of the Jardin des Plantes, Paris. IAC = Herbarium of the Iowa Agricultural College. K = Herbarium of the Royal Botanic Gardens, Kew. L = Linnaean Herbarium, Linnaean Society of Botany, London. M = Herbarium of the Missouri Botanical Garden. MU = Herbarium of the University of Missouri. NY = Herbarium of the New York Botanical Garden. P = Parry Herbarium deposited at the Iowa Agricultural College. S = Herbarium of the Botanical Museum, Stockholm. SM = Herbarium of the State Museum, Albany, New York. UC = Herbarium of the Botanical Museum of the University of Copenhagen. US = United States National Herbarium. V = Herbarium of the Natural History Museum, Vienna. Taxonomy Halenia Borckh. in Roemer, Arkiv fur Botanik l 1 : 25. 1796; Ruiz & Pavon, FL Peruv. 3 : 21, pi 2J$ } fig. 1. 1802; Endl. Gen. PL 601. 1836-40; Grisebach, Obs. Gent. 36. 1836; G. Don, Gen. 138 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 20 Hist. 4: 177. 1S38; Grisebach, Gen. & Sp. Gent. 322. 1839 Dietrich, Syn. PL 2: 91S. 1840; Hooker, Fl. Bor.-Am. 2: 67 pi. 156-6. 1840; Grisebach in DC. Prodr. 9: 128. 1845 Grisebach in Linnaea 22: 45. 1849; Weddell, Chlor. And. 2 74. 1859; Benth. & Hooker, Gen. PI. 2: 817. 1876; Hemsl Biol. Cent.-Am.Bot. 2:351. 1882 ; Baillon, Hist. PL 10:142. 1891 Gilg in Engler & Prantl, Nat. Pflanzenfam. 4 2 : 89. 1895 Conzatti, Fl. Syn. Mexico, 174. 1897; Rouy, 111. PL Eur. 17: pi. 412. 1902; Gilg in Fedde, Rep. Spec. Nov. 2:52. 1906; Robinson in Gray's Manual, ed. 7. 659. 1908; Britton & Brown, 111. Fl. 3 : 15,fig.3366. 1913;Gilg in Engl. Bot. Jahrb. 54: Beibl. 118, p. 93. 1916; Johnson, Tax. FL PL 488. fig. 340. 1931; Briquet in Candollea 4: 317. 1931 ; Rydberg, Fl. Cent. N. Am. 636. 1932. Tetragonanthus Gnielin, FL Sib. 4: 114, pi. 53. 1769; Kuntze, Rev. (Jen. PL 2: 431. 1891; Britton, Manual, 734. 1901; Small, Fl. Southeastern U. S. 931. 1913; Rydberg, Fl. Rocky Mts. 666. 1922. Swertia Linnaeus, Amoen. Acad. 2: 344. 1751; Jussieu, Gen. PL 158. 1791; Michaux, Fl. Bor.-Am. 1: 97. 1803; Humboldt, Bonpland & Kunth, Nov. Gen. & Sp. PL 3: 174. 1818; Kunth, Syn. PL 2: 266. 1823; Schlectendal //. bretricornis . 4. H. macrantha Gilg in Engl. Bot. Jahrb. 54: Beibl. 118, p. 105. 1916. Perennial herb, 5-6 cm. high, with root curved-erect, ligneous, 49 H. adpressa Allen, sp. nov. — Herba perennis, usque ad 2.5 dm. alta; radice crassa; caule plerumque solitario, erecto, tenui; foliis basalibus in rosula densa, j oblanceolatis, 1.-1.5 cm. longis, .3 cm. latis; foliis caulinis 5-6 geminis, sessilibus, lanceolatis, 1-1.5 cm. longis, .2-.3 cm. latis, acutis; inflorescentia multiflorifera cyma, pedicellis 2 cm. longis; calycis lobis lanceolatis, .4-6 cm. longis, .15 cm. latis, ob- solete 3-nerviis, medio-nervo prominenti; corolla .7-9 cm. longa, flava, tubo l /2 totae corollae longitudini parum subaequanti; corollae lobis ovatis, auriculatis, papil- latis, apiculatis; calcaribus crassis vel tenuibus, gibbis, pendulis, plus minusve divaricatis; staminibus ca. .4 cm. longis; filamentis linearibus, antheris ovalibus; capsula lanceolata; seminibus ovoideis. — Colombia: Dept. Santander, P&ramo de las Vegas, alt. 3700-3800 m., Dec. 20-21, 1026, Killip & Smith 15679 (M type, NY, US). 1933] ALLEN — THE GENUS HALENIA 149 rather elongate, covered with the blackish remains of marcescent leaves; flowering stem single, erect, internodes 4-5 cm. long, with no rosette leaves before anthesis; leaves herbaceous, acute, 3-nerved veins sunken above, prominent below; lower cauline leaves oblanceolate, gradually narrowed into a long but broad petiole, 6-7 cm. long, .5-1.5 cm. broad; upper leaves lanceolate to ovate-lanceolate, broadly sessile, about 3 cm. long; inflores- cence an apical 3-flowered cyme, solitary in axils of the upper- most euphylloid leaves; pedicels of apical flowers 3 cm. long, those of laterals 2 cm.; calyx-lobes oblanceolate, 1-1.3 cm. long, 3-nerved, acute; corolla about 1.5 cm. long, green, tube about one-third the length of the entire corolla; lobes ovate to broadly ovate, very acute, somewhat apiculate; spurs large, globose, conical, calluses about .3 cm. long at the base of the tube. Distribution : Colombia. No specimens examined, but description compiled from original publication. (type, Kalbreyer 702, BG). 5. H. Karstenii Gilg in Engl. Bot. Jahrb. 54: Beibl. 118, p. 105. 1916. Biennial or perennial herb up to 5 dm. high; root branched; stems simple, covered with leaf-bases for 7-8 cm. at intervals of less than .4 cm., internodes up to 4-5 cm. long; leaves in pairs at the nodes, sessile, lanceolate, narrowed at base, 2-5.5 cm. long, up to .9 cm. broad, 3-nerved, veins excurrent, forming a mucro; inflorescence 4-12 flowers in terminal and axillary cymes, pedicels erect, up to 4 cm. long; calyx-lobes lanceolate, .6-.8 cm. long, .2-.25 cm. broad, acuminate, 3-nerved, papillate; corolla about 1 cm. long, tube not quite one-half the length of the entire corolla; lobes ovate, subrotund, erose at apex; spurs midway up the corolla tube, small, scarcely noticeable, glandular convex depressions in the corolla; stamens about .3 cm. long, at the ori- fice of the tube; filaments linear, anthers oval, acute; capsule 15-18 cm. long, ovate; seeds ovoid, light brown, wrinkled. Distribution: paramos of Bogota. Specimens examined: Colombia: Paramo de Bogota, Karsten (V type). 6. H. subinvolucrata Gilg in Engl. Bot. Jahrb. 54: Beibl. 118, p. 99. 1916. 150 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 20 Perennial herb, about 2 dm. high; root nmltifibrous; subter- ranean stem vertical, short, thick, flowering stem erect, loosely- leafy; leaves thickly herbaceous or somewhat fleshy, 3-nerved, veins sub-parallel, deeply sunken above, prominent below; basal leaves linear-lanceolate, gradually narrowed at the base into narrow petiole, 4-5 cm. long, .4-.5 cm. broad; lower cauline leaves similar to basal leaves; upper cauline leaves broadly sessile, lanceolate or lanceolate-ovate to ovate, 2-A cm. long, .6-1 cm. broad; inflorescence terminal or axillary, almost sessile 3-flowered cymes, peduncles .3-6 cm. long; floral leaves sheathing inflorescence and longer than flowers; calyx-lobes oblong-ovate, about 1 cm. long, up to .3 cm. broad, 5-nerved, minutely hirtellous, acute; corolla about 1 cm. long, tube about one-third or less the length of the entire corolla; lobes oblong-ovate, .7 cm. long, .3-.35 cm. broad, subrotund, erose at apex; spurs scarcely prom- inent blackish maculations at the base of the tube; stamens about .2 cm. long; anthers ovate-oblong; filaments linear. Distribution: Venezuela. Specimens examined: Venezuela.: high mountains of Trujillo and Merida, alt. 1300-4900 m., 1842, Linden 437 (DH type). 7. H. inaequalis Wedd. Chlor. And. 2: 78. pi. 63 C. 1859; Gilg in Engl. Bot. Jahrb. 54: Beibl. 118, p. 99. 1916. Herb 1.5-2(-3) dm. high; stem solitary, erect, loosely leafy, with in tern odes about 4-5 cm. long, and inflorescence comprising the upper third of the stem, or stem short, leafy, with internodes equidistant, 1-2 cm. long, arising in axil of every leaf; radical leaves petiolate, lanceolate to oblanceolate, 2-3 cm. long; upper cauline leaves sessile, lanceolate to elliptic, minutely hirtellous; flowers disposed in small loose pedunculate cymules; peduncles erect or slightly nodding, 1-2 cm. long; calyx-lobes lanceolate or oblong-lanceolate, acute, margin minutely hirtellous; corolla scarcely 1 cm. long, exceeding the calyx by one-fourth its length ; lobes ovate, acute; spurs inconspicuous. Distribution: known only from Venezuela. Specimens examined: Venezuela: Paramo de Timotes, Merida, alt. 3S00 m., Sept. 4, 1921, Jahn 558 (HP, US); same locality, alt. 3600 m., Jan. 21, 1922, Jahn 839 (HP, US); Merida, alt. 3000 m., 1846, Funck & Schlim 901 (DH type). 1933] ALLEN — THE GENUS HALENIA 151 Funck & Schlim 11 48 is cited by Weddell in the original publication of the species, but Gilg has placed this number under H. viridis. 8. H. viridis (Griseb.) Gilg in Engl. Bot. Jahrb. 54: Beibl. 118, p. 100. 1916. Gentiana viridis Griseb. in Linnaea 22: 43. 1849; Wedd. Chlor. And. 2:62. 1859. H. inaequalis Wedd. Chlor. And. 2: 78. 1859; Gilg, I. c. Perennial herb, 1-3 dm. high; stem solitary, erect, thick, simple, internodes 4-5 cm. long; basal leaves in rosette, more or less coriaceous, sessile, lanceolate, subequal, up to 6 cm. long, .4 cm. broad, minutely hirtellous, 3-nerved; cauline leaves 3-4 pairs, approximately 5 cm. long; inflorescence a narrow racemi- form cyme arising in the middle of the stem, with simple 3-5- flowered cymules; pedicels erect before anthesis, cernuous after, about 2.5 cm. or less long; calyx-lobes oblong-lanceolate, minutely hirtellous, about .4-5 cm. long, nerved; corolla about 1 cm. long, greenish, tube about one-third the length of the entire co- rolla; lobes elliptic-lanceolate, rather obtuse, somewhat erose at apex; spurs inconspicuous; stamens .5 cm. long; anthers ovate; capsule oblong-lanceolate. Distribution: known only from Venezuela. Specimens examined: Venezuela: Sierra Nevada, Merida, alt. 3300 m., 1846, Funck & Schlim 1148 (DH ttpe); Laguna Mucuy, Cabeceras del Saisay, Merida, 420CH1300 m., April 19, 1930, Gehriger 92a (HP). 9. H. Schultzei 60 Gilg, n. sp. Caespitose mat-like perennial, about 3 dm. high; root tough, fibrous; stems one or more, erect, simple, slender, internodes 5-7 80 H. Schultzei Gilg, sp. nov. — Herba perennis, caespitosa, ca. 3 dm. alta; radice lenta, fibrata; caulibus 1-multis, simplicibus, tenuibus, internodiis 5-7 cm. longis; foliis basalibus in rosula densa, in petiolis longis tenuibus attenuatis, ad basin dilatatis, lineari-lanceolatis, ca. 5 cm. longis, .5 cm. latis, nerviis, acutis; foliis caulinis plerumque 2 geminis, sessilibus, linearibus, plerumque ca. 1.5 cm. longis, nerviis, acutis; inflorescentia pauco-florifera cymis terminalibus lateralibusque, pedicellis tenuibus, 1-1.5 cm. longis, ad apicem recurvatis; calycis lobis lanceolatis, ca. .6 cm. longis, acutis; corolla latissime campanulata, ca. 1.3 cm. longa, flava; corollae lobis ovatis, obtusis, marginibus erosis vel crispis; calcaribua inconspicuis. — (type, Schultze 1304, BG). [Voi-. 20 152 ANNALS OF THE MISSOURI BOTANICAL GARDEN cm. long; basal leaves in dense rosette, narrowed into long slender petioles, dilated at base, linear-lanceolate, about 5 cm. long, .5 cm. broad, nerved, acute; stem-leaves usually 2 pairs, sessile, linear, usually about 1.5 cm. long, nerved, acute; inflores- cence few-flowered in terminal and lateral cymes, pedicels slender, 1-1.5 cm. long, recurved at tip; calyx-lobes lanceolate, about .6 cm. long, acute; corolla very broadly campanulate, about 1.3 cm. long, yellow; corolla-lobes ovate, rather obtuse, erose or crisped margins ; spurs not discernible. Distribution: Colombia. No specimens examined, but description compiled from photograph, (type, Schvltze 1804, BG). 10. H. gentianoides Wedd. Chlor. And. 2 : 78. pi. 53B. 1859. Perennial herb, up to 3 dm. high; root more or less thick, ligneous; stems sterile or flowering, long, loosely leafy; basal leaves in a dense rosette, petiolate; cauline leaves 2-6 pairs, subsessile, lanceolate, acute, 3-nerved; inflorescence lateral or terminal, small racemiform cymes, peduncles more or less resu- pinate ; calyx-lobes oblong, up to .7 cm. long, papillate, acuminate, nerved tube mately one-third the length of the entire corolla; lobes ovate rotund, crenulate-erose ; spurs reduced to inconspicuous glandulai areas not visible to the naked eye; stamens approximately .4 cm anthers late. Distribution: paramos of Colombia. Specimens examined: Colomiiia: Paramo de Bogota, Karsten (V); same locality, Goudot (BG type, M photo, V); same locality, Triana 1964 (DIT, V); same locality, Jan. 17, 1854, Holton 467 (DII, G); Guasca, 1919, Ariste-,1 oseph A423 (US). 11. H. stellarioides (iilg in Engl. Bot. Jahrb. 54: Beibl. 118, p. 100. 1916. Perennial? herb, up to 3 dm. high; flowering stem erect, loosely leafy, internodes 4-7 cm. long; basal leaves lacking; cauline leaves sessile, lanceolate to linear-lanceolate, 2.5-3.5 cm. long, gradually decreasing toward the summit, .4-.5 cm. broad, acute, 3-nerved; inflorescence lateral or terminal, 5-7-flowered cymes, pedicels more or less erect, up to 1.5 cm. long; calyx-lobes lance- 1933] ALLEN — THE GENUS HALENIA 153 olate or oblong-lanceolate, about .8 cm. long, .25 cm. broad, acute, 3-nerved, or nerves obsolete; corolla about 1 cm. long, tube about one-third the length of the entire corolla ; lobes ovate- oblong, upper margin subcrenulate, acute; spurs semi-globose callous prominences at the base of the tube. Distribution: p&ramos of Colombia. No specimens examined, but description compiled from original publication and photograph, (type, Lehmann 3080, BG). 12. H. verticillata Gilg in Engl Bot. Jahrb. 54: Beibl. 118, p. 103. 1916. Annual herb, slender, up to 1 m. high; root fibrous; stem .5 cm. thick, becoming black, covered with the remains of early leaves, densely leafy at the middle or just below the middle of the stem, in- ternodes up to 8 cm. long ; basal leaves in dense whorls for a varying distance up the stem, connate, sessile, linear-lanceolate, 10 cm. or less long, .8 cm. broad, acuminate, 5-nerved, fleshy; cauline leaves in 7-8 whorls of 3, lanceolate (extreme upper subtending inflorescence, ovate-lanceolate), 5-nerved, 5-6 cm. long, gradually decreasing in length toward the summit, with corresponding increase in width; flowers numerous, disposed in 3-5 verticillate, axillary and terminal cymose clusters; pedicels erect, up to 3.5 cm. long; calyx-lobes oblong-lanceolate, papillate on under surface of veins and entire upper tip, up to 1.0 cm. long, .2-.35 cm. broad, acu- minate to acute, 5-nerved, veins parallel, becoming confluent at tip; corolla apple-green or yellowish-green, up to 1.7 cm. long, tube about one-third the length of the entire corolla; lobes broadly ovate, subrotund, erose and papillate at tip; spurs sub- globose glandular prominences at the base of the corolla. Distribution: wet pdramos about volcanos, Colombia. Specimens examined: Colombia: "Cauca am Vulkan," Sotard, 3500 m., Lehmann 6190 (BG type, F); Dept. of El Cauca, Mt. Pan de Azucar, alt. 3500-3700 m., June 16, 1922, Pennell 7052 (ANSP, G, NY, S, US). 13. H. foliosa Gilg in Engl. Bot. Jahrb. 54: Beibl. 118, p. 101. 1916. Biennial herb (?), up to 8 dm. high; root stout (?); flowering stems 1-many, erect, simple, few or no leaves at base, lower internodes 1-1.5 cm. long, upper 5-7 cm. long; numerous pairs 154 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 20 of lower cauline leaves, thickly herbaceous, gradually narrowed toward the base but dilated again at base, lanceolate, 3-5 cm. long, about 1 cm. broad, acute, 3-5 nerved; upper cauline leaves herbaceous, gradually narrowed toward the base and broadly sessile, ovate-elliptic, 3-5 cm. long, about 1 cm. broad, acute to very acute, 5-nerved, veins parallel, sunken above, prominent below; terminal and axillary, loosely arranged, many-flowered elongate cymes, comprising a thyrsoid inflorescence 10 cm. long, pedicels about 2.5 cm. long; small upper leaves shorter than the inflorescence; calyx-lobes ovate-oblong, .7-.8 cm, long, .3 cm. broad, acute or very acute, 3-nerved; corolla about 1.3 cm. long, tube about one-fourth the length of the entire corolla; spurs semi-globose protuberances at the base of the tube. Distribution: p&ramos of Col6mbia. Specimens examined: Colomiua: Dept. of Bolivar-Ant ioquia, Pjtramo de Chaquiro, alt. 3000-3200 m., Feb. 23, 1918, Pennell 4268 (NY, US), (type not seen, Stubel 276, BG, M photo). 14. H. dasyantha Gilg in Engl. Hot. Jahrb. 54: Beibl. 118, p. 103. 1916. Perennial herb, (1.5— )3— 7 dm. high ; root thick ; stem erect, simple, more than .5 cm. thick, fleshy, brown in dried specimens, faintly striate, covered with remains of marcescent leaves, 2-5 internodes up to 10 cm. long; basal leaves in a dense rosette, broadly elliptic, up to 10 cm. long, nearly 2 cm. broad, acute, 5-nerved; 3-5 pairs of cauline leaves, sessile, dilated at base, oblong-lanceolate, 4-10 cm. long, their length gradually decreasing toward the summit, 1-2 cm. broad, acute, 5-nerved; inflorescence axillary and ter- minal, many-flowered racemose cymes, usually 10 cm. long; pedicels erect, for most part up to 3.5 cm. long; calyx-lobes ovate to ovate-lanceolate, papillate, up to 1 cm. long, .4-.5 cm. broad, acute to abruptly acuminate, 3-nerved; squamellae frequently scale-like lobed bodies; corolla 1.5-2 cm. long, pale greenish- yellow, tube nearly equal in length to the entire corolla; lobes ovate, subrotund, erose at apex, somewhat papillate; spurs small subglobose prominences at the base of the corolla-lobes, almost obscured by the calyx; stamens approximately .4 cm. long, attached at the summit of the tube; filaments linear, anthers ovate-oval; stigma reflexed; capsule up to 2 cm. long; seeds elliptical, reticulate. 1933] ALLEN — THE GENUS HALENIA 155 Distribution: moist grassy paramos, or dry open woods, Colombia. Specimens examined: Colombia: Dept. of Caldas, Paramo del Quindio, alt. 3700-4200 m., Aug. 15-20, 1922, Pennell & Hazen 9997 (ANSP, NY, US); Dept. of Tolima, Paramo de Ruiz, alt. 3500-3800 m., Dec. 16-17, 1917, Pennell 3001 (NY, M, US); Dept. of Cauca, Paramo de Buena Vista, Huila group, Central Cordillera, alt. 3000-3600 m., Jan. 1906, Pittier 1111 (US), (type not seen, Lehmann 2065, BG, M photo.). This species is very similar to H. elata but is, on the whole, a larger and coarser plant. Since the material is scanty and the geographical location different, H. dasyantha has for the present been maintained as a distinct species. 15. H. elata Wedd. Chlor. And. 2: 78. 1859. Perennial, up to 5 dm. high; stem thick, coarse, erect, loosely leafy; numerous basal leaves in dense rosette, broadly lanceolate, attenuate into petiole, dilated at base, 7-10 cm. long, approxi- mately 1 cm. broad, 5-nerved, acuminate; cauline leaves more or less sessile, shorter, lanceolate, acute; inflorescence numerous loose terminal and lateral cymes, peduncles up to 3 cm. long; calyx-lobes oblong-lanceolate to ovate-lanceolate, .7-.8 cm. long, 3-nerved, papillate, acute; corolla hardly one-fourth more than the length of the calyx, tube slightly more than one-half the length of the entire corolla ; lobes ovate, obtuse ; spurs inconspicu- ous tubercules at the extreme base of the corolla; stamens .6 cm. long; anthers narrowly ovate, filaments linear. Distribution: in the Sierra Nevada of Santa Marta, Venezuela. Specimens examined: Venezuela: Sierra Nevada de Santa Marta, Caracas, 1844, Funck 415 (DH type). 16. H. Tolimae Gilg in Engl. Bot. Jahrb. 54: Beibl. 118, p. 101. 1916. Perennial herb, 2.5-3 dm. high; root thick; root-stalk thick, densely covered with remains of marcescent leaves; one to few flowering stems, thick, erect, simple for the most part, very narrowly winged, internodes 3-6 cm. long; basal leaves arranged in rosette, slightly narrowed into long broad petioles, dilated at the base, lanceolate, 5-8 cm. long, up to .5-.6 cm. broad, 3-5- nerved, acute; stem-leaves 2-3 at a node, sessile, broadly lanceo- late, 4-6 cm. long, .6-.7 cm. broad, their size gradually de- creasing toward the summit, 3-5 subparallel veins, sunken above, [Vol. 20 15G ANNALS OF THE MISSOURI BOTANICAL GARDEN prominent below, reticulate; inflorescence few-flowered (5-7) cymes, axillary and terminal; pedicels more or less resupinate, up to 3 cm. long, the central one longer; calyx-lobes ovate to ovate- oblong, papillate, .7-.9 cm. long, .25-.35 cm. broad, subacuminate, 3-nervcd; corolla up to 1.3 cm. long, light greenish-yellow, tube slightly less than one-third the length of the entire corolla; lobes ovate, erose and papillate at the tip; spurs glandular subglobose protuberances at the base of the tube, obscured by the calyx; stamens approximately .4 cm. long; filaments linear, anthers ovate; capsule lanceolate, apiculate, 2 cm. long; seed ovoid- elliptical, very minutely reticulate, pale tan. Distribution: grassy paramos of Colombia. Specimens examined: Colombia: Dept. of Caldas, I'aramo del Quindio, alt. 4100-4400 m., Aug. 15-20, 1922, Peiinell & Hazen 9841 (ANSP, G, NY, US); (type not seen, Stilbcl 228, BG, M photo.); bare loam slopes below snow, same locality and date, alt. 4300-4500 m., Pennell A Hazcn 9894 (ANSP, NY, S, US). This last-cited specimen has leaves broader than those of the type, but is similar otherwise. //. Tolimae appears to be closely related to H. data. 17. H. hygrophila Gilg in Engl. Bot. Jahrb. 54: Beibl. 118, p. 102. 1910. Biennial herb, about 6 dm. high; root ligneous; subterranean root-stalk covered with darkened leaf-bases; flowering stem fleshy, erect, simple, 1 cm. thick, faintly winged, internodes up to 8 cm. long; 30-40 basal leaves in a dense rosette, linear, up to 12 cm. long, .5-.9 cm. broad, acute, 5-nerved; cauline leaves yellow-green, 3-4 pairs, the lower lanceolate, the upper sub- tending the inflorescences, ovate-lanceolate, sessile, 5-7 cm. long, gradually decreasing in size toward the summit, approxi- mately .8-1.2 cm. broad, 7-nerved, parallel veins confluent at tip; inflorescence terminal and axillary in upper stem-leaves, in dense many-flowered cymes, giving spicate appearance, pedicels erect, up to 3 cm. long; calyx-lobes ovate-oblong to ovate, papillate, .9-1.0 cm. long, narrowly acute, 5-nerved, reticulate; corolla ud to 1.5 cm. lone, vellowish-ereen, tube approximated third the length of extreme base of the corolla lobes, obscured by calyx; stamens approximately .7 cm. lone, attached at the orifice of the tube; filaments linear. 1933] ALLEN — THE GENUS HALENIA 157 anthers broadly oblong, apiculate; capsule lanceolate; seeds elliptic, pale tan, reticulate. Distribution: pdramos of Andes, Colombia. Specimens examined: Colombia: Paramo of Guanacas, Central Andes of Popayan, 3000-3600 m., Lehmann 7860 (F, G, BG type, M photo, US). 18. H. parallela 51 Allen, n. sp. Perennial herb, up to 6 dm. high; root-stalk thick; flowering stem single, erect, simple, or branched above, striate, lower inter- nodes 1-1.5 cm. long, upper 4-6 cm. long; numerous basal leaves in rosette, almost sheathing, lanceolate, 3-5-nerved, about 9 cm. long, 1 cm. broad; cauline leaves about twelve pairs, more remote toward the apex, sessile, lanceolate to elliptic, 4-6 cm. long, gradually decreasing toward the summit, approximately 1 cm. broad, 3-5 subparallel veins sunken above, prominent below; inflorescence 1 or more many-flowered axillary and terminal cymes, pedicels more or less erect, up to 3.5 cm. long, the lateral somewhat shorter than the terminal; calyx-lobes ovate, papillate, .7-9 cm. long, .4 cm. broad, attenuately acute, prominently 5-7-nerved, veins subparallel; corolla about 1.3 cm. long, probably greenish?, tube about one-fourth the length of the entire corolla; lobes broadly ovate, slightly papillate, crisped toward the tip, abruptly acuminate ; spurs large globose protuberances at the base of the tube, obscured by calyx; stamens nearly .5 cm. long, attached at the orifice of the tube; filaments linear, anthers oblong; capsule lanceolate. 61 H. parallela Allen, sp. nov. — Herba perennis, usque ad 6 dm. alta; radice crassa; caule florifero solitario, erecto, simplice vel supra ramoso, striato; internodiis inferioribus 1-1.5 cm. longis, superioribus 4-6 cm. longis; foliis basalibus multis, in rosula densa, fere vaginantibus, lanceolatis, 3-5-nerviis, ca. 9 cm. longis, 1 cm. latis; foliis caulinis ca. duodecem geminis, ad apicem remotioribus, sessilibus, lanceo- latis vel ellipticis, 4-6 cm. longis, sensim sursum decrescentibus, ca. 1 cm. latis, 3-5-nerviis, nervis subparallelis, supra immersis, infra prominentibus; inflor- escentia cymis 1- vel pluro-floriferis axillaribus et terminalibus; pedicellis plus minusve erectis, usque ad 3.5 cm. longis, lateralibus terminalibus brevioribus; calycis lobis ovatis, papillatis, .7-9 cm. longis, .4 cm. latis, attenuate acutis, pro- minente 5-7-nerviis, nervis subparallelis; corolla ca. 1.3 cm. longa, forte viridula, tubo ca. l /i totae corollae longitudini adaequanti; lobis late ovatis, parum papillatis, ad apicem crispis abrupte acuminatis; calcaribus magnis globosis gibbis ad basin tubi calyce obscuratis; staminibus ca. .5 cm. longis, tubi summo adjunctis; fila Venezuela: Pdramo de TYPE [Vol. 20 158 ANNALS OF THE MISSOURI BOTANICAL GARDEN Distribution: Venezuela. Specimens examined: Venezuela: IViramo de La Negra, M6rida, Dec. 1927, Gutzmller 82 (HP type, US). Species near H. foliosa and H. hygrophila. 19. H. major Wedd. Chlor. And. 2: 79. 1859. Annual probably, up to 6 (?) dm. high; (root not seen); stem erect, simple below, branched above, loosely leafy, very narrowly winged and striate; leaves sessile, subconnate, broadly elliptic, 2-7 cm. long, .5-2 cm. broad, somewhat abruptly acuminate, 3-5-nerved; inflorescence terminal and axillary 3-6-flowered sub- umbelliform cymes, pedicels 4 cm. long, usually erect, frequently nodding at apex; calyx-lobes oblanceolate to subspatulate, up to .8 cm. long, .25 cm. wide, 3-nerved, midvein very prominent; corolla 1.0-1.3 cm. long, tube almost one-half the length of the corolla; lobes ovate, erose at apex; spurs small upcurved conical protrusions almost at the orifice of the tube; stamens approxi- mately .5 cm. long, at the orifice of the tube; filaments linear; anthers not seen; capsule ovate, 1.3-1.6 cm. long; seeds oval, brown-black, wrinkled. Distribution: shrub zone, mountain bases, Colombia. Specimens examined: Colombia: Dept. of El Cauca, Mt. Pan de Azucar, alt. 3300-3600 m., June 16, 1922, Pennell 7084 (ANSP, NY, US); Dept. of Cundinamarca, Sibate, alt. 2700-2800 m., Oct. 13-15, 1917, Pennell 2438 (M, US); Dept. of Caldas, Cerro Tatama, alt. 3400-3700 Goudot Suction 2. IIaleniastrum KEY TO NORTH AMERICAN SPECIES AND VARIETIES 1. Spurs spreading to ascending. 2. Plant less than 2.5 dm. high. 3. Biennial; flowers white; distribution Mexico 20. 11. crassiuscula 3. Annual; flowers purple; distribution Canada . 28a. II. dejh.ra var. Brentoniana 2. Plant more than 2.5 dm. high. 3. Spurs, if present, about Y 2 the length of the corolla 21. H. Pringlei 3. Spurs less than Yi the length of the corolla. 4. Annuals; leaves mostly cauline. 5. Stem-leaves linear 22. 11. rrcurva 5. Stem-leaves lanceolate to ovate 23. II. deflexa 4. Perennials; leaves mostly radical. 5. Strict, many-flowered, spike-like inflorescence. 6. Stem erect 24. H. rhyacophUa 6. Stem procumbent 24a. H. rhyacophUa var. procumbens 1933] ALLEN THE GENUS HALENIA 159 5. Loose, broad, few-flowered inflorescence 24b. H. rhyacophila var. macropoda 1. Spurs pendulous to incurved. 2. Leaves not apiculate or very rarely so; calyx-segments not mucronulate. 3. Basal rosette absent; leaves mostly cauline. 4. Habit erect; leaves less than 3 cm. long. 5. Leaves linear 25. H. Palmeri 5. Leaves lanceolate or ovate. 6. Calyx-lobes mostly obtuse, appressed 26. H. Conzattii 6. Calyx-lobes acute, reflexed 27. H. Schiedeana 4. Habit prostrate; leaves 5-12 cm. long 28. H. caleoides 3. Basal leaves present, cauline few or none. 4. Spurs incurved, Y2 or less than Y2 the length of the corolla. 5. Sterile branches present; leaves numerous 29. H. platyphylla 5. Sterile branches absent; leaves less than 15. 6. Flowers 1.5 cm. or less long. 7. Flowers less than 1 cm. long; spurs rudimentary. .30. H. nudicaulis 7. Flowers more than 1 cm. long; spurs %- l A the length of the corolla 31. H. plantaginea 6. Flowers more than 1.5 cm. long. . .3 la. H. plantaginea f. grandiflora 4. Spurs spreading, about K the length of the corolla. 5. Stems always erect. 6. Leaves linear 32. H. Shannonii 6. Leaves elliptical 32 a. H. Shannonii f . compacta 5. Stems more or less decumbent S3. H. decumbens 2. Leaves conspicuously apiculate; calyx-segments mucronulate. 3. Plant more than 2 dm. high 34. H. guatemalensis 3. Plant less than 2 dm. high 34a. H. guatemalensis var. latifolia 20. H. crassiuscula Robinson & Seaton in Proc. Am. Acad. 28: 113. 1893. Small caespitose biennial of dense habit, slightly fleshy ; stems erect, 0.4-1.0 dm. high, narrowly winged, much branched; radical leaves broadly oblanceolate to elliptic, 2 cm. long, obtuse, attenuate into long petiole nearly equalling the blade, 3-nerved; cauline leaves 1-3 pairs, narrowly oblanceolate to oblong, nar- rowed at the base; inflorescence dense compact umbellate cyme; flowers terminal or axillary, pedicellate, after anthesis slightly nodding, not at all resupinate; calyx-segments lanceolate to oblanceolate, .45-.6 cm. long, obtuse, papillate, 3-nerved; corolla white, up to 1.5 cm. long, about .5 cm. broad at base, tube .35-.4 cm. long; lobes oblong-elliptic, acute; spurs .4 cm. long, arising slightly below the midpoint of the tube, slender, spreading, and curved upwards; stamens .25 cm. long, anthers ovate; filaments linear; capsule lanceolate, frequently subfalcate, acute, 1.4 cm. long; seeds globose, light yellow-brown, granular. 160 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 20 Distribution: bare alpine summits, Mexico. Specimens examined: Mexico: Nevado de Toluca, bare alpine summits, alt. 3500 m., Sept. 2, 1892, Pringle 4229 (ANSP, BG, BM, B, CAS, C, DH, F, G type, IAC, K, M, NY, S, US, V); fartacvihuatl, wet meadows, alt. 3000-3250 m., March-July 1903, Purpus 318 (CAS, M, US); Popocatepetl, Aug. 7-8, 1901, Rose & Hay 5999 (US). 21. H. Pringlei Robinson & Seat on in Proc. Am. Acad, 28: 113. 1S93. Halenia Candida Ramirez in Inform. Secret. Foment. Mexico (Excurs. Mont. Ajusco), 34. 1895; Estud. Hist. Nat. 102. 1904. Biennial, of glaucous aspect; root thick, tough, ligneous; stem usually solitary, occasionally caespitose, simple or nearly so, scape-like, slender, erect, 1-2.5 dm. high; leaves less than 3 cm. long, radical leaves elliptic to narrowly oblanceolate, faintly 3- nerved, attenuate below into slender petioles, usually as long as the leaf-blade and persistent; cauline leaves 1-2 pairs, sessile, short, sublinear, 1.5-3 cm. long, about .3 cm. broad; flowers disposed in terminal, or occasionally lateral, few-flowered um- belliforni cymes, pedicels less than 2 cm. long, usually about .8 cm.; calyx-lobes oblong-spatulate, .3-5 cm. long, acuminate, 3- nerved; corolla white, .8-1.5 cm. long, spurless in the majority of cases; when spurs present, slender, spreading, and curved- ascending, 1.5-1.8 cm. from tip to tip, with prominent veins and glandular tips; corolla-tube up to .2 cm. long; corolla-lobes elliptic, acuminate; stamens .2-35 cm. long; anthers narrowly ovate, minutely papillate, filaments linear, slightly uncinate; capsule lanceolate, acute, exserted; seeds subglobose, yellow- brown, granular. Distribution: springy meadows of central and south Mexico. Specimens examined: Mexico: springy alpine meadows, Sierra de las Cruces, alt. 2450 m. f Aug. 28, 1904, Pringle 13121 (BG, C, UC, G, K, US); same locality, Aug. 23, 1892, Pringle 4209 (ANSP, C, CAS, DH, F, G type, IAC, K, M, NY, S, US, V); same locality, June 1895, Altamirano 90S (US). South Mexico: without locality, 1920-21, Revoke 36 (BG). The habit of this species is very similar to that of H. nudicaulis. The Pringle specimens cited above were collected in August, and the plants are smaller and grow less luxuriantly than the single specimen collected in June by Altamirano. The specimens collected later in the season verv rarelv oossess sours, while the 1933] ALLEN — THE GENUS HALENIA 161 earlier plants show a distinctly spurred corolla. This condition is shown in other species to a somewhat less extent, and is in all probability traceable to variation in environmental conditions. 22. H. recurva (Sm.) Allen, n. comb. Swertia recurva Smith in Rees, Cyclopedia 34: sub Swertia. 1819. Halenia Rothrockii Gray in Proc. Am. Acad. 11: 84. 1876; Rothrock, Rept. Wheeler Exped. 195, pi 21. 1878; Hemsl. Biol. Cent.-Am. Bot. 2 : 353. 1882. Tetragonanthus Rothrockii Heller, Cat. N. Am. PI. 6. 1898, and ed. 2, 16. 1900. Annual, 2.5-5 dm. high; stem simple, often branched above; basal leaves less than 3.5 cm. long, .6 cm. broad, elliptic-lanceolate to spatulate; cauline leaves remote, lance-linear, 1.5-4 cm. long, about .35 cm. broad, obscurely 3-nerved, midrib prominent below; inflorescence a loosely flowered subumbellate cyme; flowers on slender pedicels, .5-3 cm. long, often in sevens; calyx lobes lanceolate, elongate-acute, up to .6 cm. long, uninerviate, papillate; corolla bright-yellow, about 1-1.2 cm. long, tube less than one-half the length of the entire corolla ; corolla-lobes ovate, subacuminate, delicately veined, papillate; spurs curved, hori- zontal or ascending, up to 1.6 cm. from tip to tip; anthers broadly oblong, mucronate, papillate; filaments slightly obovate; capsule ovate-lanceolate; seeds yellow-brown, subglobose-ovoid, granular. Distribution : southern United States and Mexico. Specimens examined: United States: Arizona: Mt. Graham, alt. 2250 m., Sept. 1874, Rothrock 733 (ANSP, F, IAC, M, US); same locality, Aug. 1874, Rothrock (G, NY); Chiricahua Mts., Sept. 22. 1931, Jones 28603 (M); Barfoot Park, Chiricahua Mts., alt. 2000-2050 m., rolling andesitic, recently lumbered pine land, Sept. 8, 1906, Blumer 1359 (BG, D, G, K, M, NY, US, V); same locality, alt. 2480 m., Sept. 22-23, 1914, Eggleston 10774 (US); Apache Pass, Chiricahua Mts., Sept. 1881, Lemmon & Lemmon (CAS) ; Hermitage, Chiricahua Mts., Sept. 1881, Lemmon & Lemmon (CAS); Rucker Valley, Chiricahua Mts., Sept. 1881, collector unknown 1874 (CAS, DH, F, SM); White Mts., Aug. 1873, Lour (F); summit of White Mts. (Springerville-Fort Apache Road), Apache Reservation, alt. 2270-2880 m., Aug. 29, 1919, Eggleston 15781 (F); Riverside Ranger Station, Greer, Apache Forest, alt. 2700 m., Aug. 24, 1920, Eggleston 17137 (NY, US); grassy flats near Brinkley's Ranch, White Mts., Aug. 5, 1915, Ellis 20 (US); Riggs Flat, Pinaleno Mts., alt. 2000 m., Sept. 23, 1917, Shreve 5373 (G); Columbia Trail, Pinaleno Mts., alt. 2500 m., Sept. 13, 1914, Shreve 4812 (CAS, US). 162 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 20 New Mexico: Mogollon Mta., on or near the west fork of the Gila River, Socorro Co., alt. 2125 in., Aug. 14, 1903, Metcalfe 601 (M, NY); swampy ground, divide of Mogollon Mts., Sept. 7, 1881, Rmby 264 (ANSP, BM, CAS, F, K, M, NY, US); same locality, Aug. 1881, Rusby (IAC, NY). Mexico: Chihuahua: Mt. Mohinora, Sept. 1, 1898, Nelson 4868 (G, US); cool slopes, Sierra Madre, alt. 1750-2375 m., Sept. 27, 1888, Pringle 1663 (BG, BM,B, CAS, DH, M, NY, S, V); same locality, Sept. 24, 1887, Pringle 1329 (ANSP, BG, C, G, K, NY, US); Meadow Valley, Sierra Madre Mts., alt. 1750 m., Sept. 17, 1903, Jones (S); Sierra Madres, near Colonia Garcia, alt. 2000 m., Sept. 6, 1899, Tovmsend & Barber 309 (BB, F, DH, G, M, NY, US); Escalon, Mutis (L). Coaiiuila: Sierra Madre, 40 in. south of Saltillo, July 1880, Palmer 839 (ANSP, G, K, US). Duran<;o: Barranca, below Sandia Station, alt. 1625 m., Oct. 13, 1905, Pringle 13588 (G, S, UC, US); Sierra de Candela, alt. 3000 m., Aug. 27, 1903, Endlich 53 (BG). Jalisco: Sierra de Tequila, alt. 2000 m., July 5, 1893, Pringle 5465 (G). The name Swertia recurva Smith was given to the specimen collected by Mutis and sent to Linnaeus, now preserved in the herbarium of the Linnaean Society of London. The description published in Rees' 'Cyclopedia' was inadequate, and later the specific name recurva was placed under deflexa, the well-known northern species. Since few of the succeeding monographers ever saw the original Swertia recurva Sm., it is not strange that the error persisted. Over fifty years later, Gray described Halenia Rothrockii as a new species. A careful examination of both specimens reveals the fact that they are identical, in which event the correct specific name is recurva. 23. H. deflexa (Sm.) Griseb. Gen. & Sp. Gent. 324. 1839; Hook. Fl. Bor.-Am. 2: 67. pi. 155. 1840; Dietrich, Syn. PI. 2: 918. 1S40; Torr. Nat. Hist. N. Y. 2 2 : 110. 1843; Robinson in Gray, Man. ed. 7, 659. 1908; Johnson, Tax. Fl. PI. 488. fig. 840. 1931; Louis-Marie, Fl. Man. Prov. Queb. 214, pi. 68, fig. 11. 1931. Swertia deflexa Smith in Rees, Cycl. 34: sub Swertia. 1819. S. corniculata Michx. Fl. Bor.-Am. 1: 97. 1803. S. americana Spreng. Syst. 1: 661. 1825. S. Michauxiana G. Don, Gen. Hist. 4: 177. 1838; Schl. & Cham, in Linnaea 5: 122. 1830. Halenia Michauxiana G. Don, Gen. Hist. 4: 177. 1838. H. heterantha Griseb. Gen. & Sp. Gent. 325. 1839. H. rejlexa Griseb. in DC. Prodr. 9: 135. 1845, sphalm. 1933] ALLEN — THE GENUS HALENIA 163 Tetragonanthus deflexus Kuntze, Rev. Gen. PL 2: 431. 1891; Heller, Cat. N. Am. PI. 6. 1898, and ed. 2, 161. 1900; Britt. Man. 734. 1901 ; Britt. & Brown, 111. Fl. 3 : 15, fig. 8865. 1913; Small, Fl. Southeastern U. S. 931. 1913. T. heteranthus Heller, Muhlenbergia 1:2. 1900. T. heterantherus Heller, Cat. N. Am. PI. ed. 2, 161. 1900, sphalm. T. deflexus heteranthus Britt. Man. 735. 1901. Annual, 1-9 dm. high; stem simple or branched above, quad- rangular; leaves 3-5-nerved, basal oblong-spatulate, 1-2 cm. long, petiolate; cauline leaves oblong-lanceolate to ovate, acu- minate, 1-5 cm. long, .5-2 cm. broad; internodes 6-8 cm. long; flowers disposed in a terminal or axillary, loose umbelliform verticillate cyme; calyx .4-.8 cm. long, segments ovate-lanceolate, acuminate, papillate; corolla purple, .8-1.4 cm. long, lobes lanceo- late to ovate, acute, papillate, tube about equalling the limb ; spurs .3-.5 cm. long, slender, cylindrical, obtuse, curved-spreading, deflexed at apex, glandular, frequently lacking in lower flowers or in flowers blooming late in the season ; stamens slightly uncinate ; anthers ovate, filaments linear; capsule lanceolate; seeds oblong- ovoid, greenish-brown, granular. Distribution: cool damp woods, from Labrador to New York, west to British Columbia and Montana; also in central Mexico. Specimens examined: (?) Labrador: Caribou Island, 1870, Macfarlane (BB); same locality, Martin (G); without locality, Rothrock (F). Newfoundland: calcareous rocks and talus, entrance to Port Saunders Harbor, Ingornachoix Bay, Aug. 1, 1910, Fernald, Wiegand & Kittredge 3911 (G); Chimney Cove, Aug. 16, 1896, Waghorne (DH, G, M); without locality, Banks (G); Brenton (K). Nova Scotia: hills between northeast Margaree and Grand Etang, Cape Breton, Aug. 13, 1906, Robinson 38 % (NY). New Brunswick: Fredericton, Aug. 1881, Bailey (US); Drury's Cove, St. Johns, Aug. 18, 1873, Boott (G); Falls, Aroostook River, Aug. 17, 1901, Churchill (M); Charlo, Restigouche, July 30, 1894, Fowler (US); Sugar Loaf, Restigouche, Bass River, Aug. 3, 4, 1873, Fowler (M); same locality and collector, Aug. 1, 1882 (F); St. Johns, July 12, 1877, Fowler (ANSP); Eel River, York Co., July 20, 1882, Hay (ANSP); open woods, Connors, July 20, 1908, Mackenzie 3618 (M, NY, US); dry fields, Rothesay, St. Johns, Maihew (BG); Saint Francis Parish, July 29, 1900, Williams (CAS, G). Quebec: St. Anne des Monts, Gasp6 Co., Aug. 16, 1881, Allen (NY); wet woods, Riviere du Loup, Canby (F); woods, Lake Memphremagog, July 22, 1902, Churchill (G); wet woods about Georgeville, Lake Memphremagog, Aug. 1, 2, 20, 1914, 164 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 20 Churchill (F, G, K, M, NY, US); Carleion, Bonaventure Co., July 23, 24, 27, 1904, Collins, Fernald & Pease 4261 (G); cool wooded banks, between Bald6 and the Baie des Chalmrs, Bonaventure River, Bonaventure Co., Aug. 5, 6, 8, 1904, Collins, Fernald A Pease (G); Rivi&re du Loup, Aug. 1902, Eggleston 3051 (DH, K, M, NY, S, US); St. Anne des Monls, July 15, 1923, Fames (SM); slaty soil, Rimouski, July 18, 1907, Fernald 1151 (G); alluvial wooded banks, Riviere Ste. Anne des Monts, Gasp6 Co., July 16, 1906, Fernald & Collins 244 (K, NY, UC, US); wood-road along Riviere Cap Chat, Matane Co., Aug. 18, 1923, Fernald, Dodge & Smith 25986 (G); Matane, near the St. Lawrence, Gasp6, Aug. 6, 1904, Forbes (G); Little Metis, Aug. 7, 1906, Fowler (G); Anticosti, Aug. 1, 1861, Hyatt, Shakr A Verrill (G); Riviere du Loup, Aug. 15, 1892, Kennedy (ANSP, G); Mt. Albert, Gasp6, Aug. 1882, Macoun (NY); wrt rocks, Salt Lake, Anticosti, Aug. 11, 1883, Macoun (BM); Anse k Persil, Rividre du Loup, July 1913, M arie-V ictorin 28 (G, NY, SM, US); Baie Girard (Lake Temiscarning), Abitibi, June 27, 1918, M arie-Y ictorin 8349 (US); "Lac Sale: dans la prairie naturelle prds de la maison du garde," Anticosti, July 23, 1927, Marie- Victorin & Holland-Germain 27159 (G)j "RiviSre Vaureal: talus calcaires," Anticosti, July 31, 1925, Marie-V ictorin, Rolland-Germain & Louis-Marie 21076 (G); "Le long de la Riviere Sainte Anne des Monts; & 10 milles de Tembouchure, Gasp6sie," Aug. 3, 1923, Marie-V ictorin, Brunei, Rolland-Germain & Rousseau 17663 (G); Notre Dame de Lac, Temiscouata Co., July 9, 1903, Moore 1211 (G); same locality, July 30, 1887, Northrup 69 (NY); George ville, July 28, 1902, Pease 1081 (G); abun- dant along shores of Lake Memphremagog, Georgeville, July 31, 1902, Pease 1082 (G); Notre Dame du Lac, Temiscouata Co., July 9, 1903, Pease 2384 (G); Georgeville, July 31, 1902, Pease 2910 (G); Riviere du Loup, St. Lawrence, 1860, Pickering (G); Lower St. Lawrence, Aug. 6, 1879, Pringle (US); shore of St. Lawrence, Temiscouata, Aug. 7, 1S79, Pringle (F, G, IAC, M, US); banks of Grand River, Gasp6 Co., June 20— July 10, 1903, Richards (G); "Cap k l'original: clairkVe; dans un bois de coniferes, Comt<5 de Rimouski, Bic," July 19, 1927, Rousseau 30711 (G, M); "Cap aux Cor- beaux: dans un bois de coniferes; sur le conglomerat. Bic, Comt6 de Rimouski/' July 14, 1927, Rousseau 26646 (M); "He Bayfield (Sandy Island), Archipel de St. Augustin," Labrador Peninsula, Saguenay Co., July 21, 1915, St. John 90688 (G); roadside, Lac du Saummi, Matane Co., Aug. 15, 1923, Svenscn & Fassett 2096 (G, SM); Anticosti, Verrill (F); damp grassy open meadow, Pointe Nouvelle, Hope Township, Bonaventure Co., July 30, 1902, Williatns & Fernald (G); Bic, July 16, 1910, Williamson 1422 (ANSP, NY). Ontario: banks of the Maitland River, 1836 Goderiels (DH); Moose Factory, Hudson's Bay, July 1, 1SXI, Haydon (K); same locality, 1880, Haydon (K); between Moose Factory and Rupert's House, southern end of Hudson's Bay, June 12, 1860, Drexter ((!); Sand River, Aug. 24, 1928, Hetnburger (COP); Pic River, Lake Superior, Loring (G); Lake Huron, Aug. 3, 1871, Macoun 2239 (DH); Whitefish Island, Lake Huron, Aug. 28, 1901, Macoun 300 (NY); "Lake Region and Ontario," July 29, 1874, Macoun 1191 (K); damp woods, Lake Niphigon, July 10, 1884, Macoun (BM); Salt, July 29, 1891, Morton (D); damp soil, Gray (mile 229 of Algoma Central Ry.), June 23, 1921, Pease 18080 (G); shaded bank, Burnt Rock Pool, Agawa R., June 21, 1921, Pease 18058 (G); moss-grown fissures of Laurentian rocks along Onaman River, Thunder Bay District, 1912, Pulling (G); Minaki, July 25, 1915, Thompson 31 (M); swamp, New Hanbury, Aug. 14, 1899, Umbach (BG, US); vicinity of Fort William, dry banks, Aug. 5, 1912, Williamson 2090 (ANSP); same locality, July 15, 1869, Macoun (K). 1933] ALLEN — THE GENUS HALENIA 165 Maine: Penobscot River, 1836, Bailey (NY); Katahdin, woods near Mountain, Blake (C, F) ; swamp woods, Chute (F) ; gravelly thicket, Boundary Lake, St. Francis River Valley, Aroostook Co., Aug. 12, 1902, Eggleston & Fernald (G); Brookline (Naskeag Point), Faxon (G); open woods, Fort Kent, Aug. 4, 1907, Fellows (US); moist banks, Aroostook Co., Aug. 25, 1893, Fernald (DH); wooded gravelly river- bank, Island Falls, valley of Mattawamkeag, Aroostook Co., Sept. 6, 1897, Fernald (G); wooded river-bank, Van Buren, Aroostook Co., Sept. 18, 1900, Fernald (G); damp, wooded slope, Hampden, Penobscot Co., Sept. 8, 1916, Fernald & Long 14392 (ANSP); argillaceous ledges, Old Town, July 27, 1916, Fernald & Long 14390 (ANSP, F, SM, US); damp, gravelly woods, Houlton, Aroostook Co., Aug. 26, 1897, Fernald (G); low woods, Orono, July 29, 1890, Fernald (ANSP, NY); moist banks, along - St. Johns River, St. Francis, Aug. 25, 1893, Fernald 87 (ANSP, C, CAS, G, K, NY, M, US); Bangor, Hallowell (BB, IAC); in woods on banks of Penobscot, Oldtowne, 1828, Oakes (G); banks of the Wassataquoik River, 1847, Porter (M); Seven Islands, Township 13, Ranges 14-15, river-bank, July 25, 1917, St. John & Nichols 2449 (NY, US); banks of Wassataquoik, Aug. 1847, Thurber (F, G, NY); roadside ledge, Frenchville, Aug. 12, 1901, Williams (G); Ashland, Fort Kent Road, "Winterville," Aug. 9, 1901, Williams (G); on Allagash River, at "Eliza Hole," Aroostook Co., July 28, 1900 Williams (G); in loam, borders of spruce woods, Portage Lake, Aug. 9, 1901, Williams, Robinson & Fernald 58 (ANSP, BG, B, CAS, D, DH, F, G, IAC, K, M, NY, SM, UC, US); banks of the Wassataquoik, common on the Penobscot and its tributaries, Aug. 1847, Young (G, NY, UC). New Hampshire: open pastures, Lombard Hill, Colebrook, Coos Co., July 20, 1917, Fernald & Pease 16624 (G). Vermont: Charlotte, July 28, 1881, Hosford (US); same locality and collector, Aug. 3, 1879 (C, F, G, IAC, M); same locality and collector, Aug. 13, 1878 (COP, G); sandy woods, Salem Lake, Derby, Sept. 3, 1931, Pinkerton & Allen (M); maple woods, West Woodstock, July 30, 1928, Kittredge (M). Massachusetts: banks of Manhan River, Southampton, 1830, Chapman (M, NY). New York: Trenton Falls, Aug. 9, 1883, Haberer (CAS, US); same locality, Aug. 18, 1902, Haberer 601 (G, SM); Fairfield, Herkimer Co., Hadley (SM); without locality or date, Hadley 1 (NY); Sylvan Beach, July 20, 1914, Home 5643 (SM); banks of the Hudson River, North Creek, Warren Co., Sept. 29, 1927, House 15688 (G, SM); Cocheton, Aug. 1, 1887, Poggenburg (NY); Trenton Falls, Aug. 18, 1902, Peck (SM); Trenton Falls, Aug. 29, 1868, Schaffer (ANSP); Cocheton, July 1887, Schrenk (NY, SM). Michigan: St. Helena Island in northern Lake Michigan, July 19, 1886, Arnold (BB); Mackinac, July 17, 1881, Boyce (IAC); without locality, Aug. 28, 1892, Dodge 352 (?) (M); near Port Huron, Aug. 16, 1892, Dodge (BB, F, G, US); same locality, Aug. 16, 1895, Dodge (US); Keweenaw Co., July 4, 1888, Farwell 249 (UC); low grounds, Keweenaw Co., Aug. 1890, Farwell 770 (G); Big Stone Bay, Emmet Co., piney-aspen woods, July 31, 1925, Gates 14156 (S); weeds in trail in Thuja bog, Reese's, Douglas Lake, Cheboygan Co., June-Aug. 1917, Gates & 9 Gates 10716 (F, M); same locality, Aug. 12, 1916, Gates & Gates 9768 (BB); Cedar Swamp, Boyne Falls, July 27, 1878, Hill 179 (F); Isle Royale, July 21, 1889, Holway (IAC, NY); Cedar Swamp, Cheboygan, Aug. 20, 1890, Kofoid (G, M); moist soil in wet places, Grayling, July 28, 30, 1903, Mell & Knopf (M); Mackinac Island, July 28, 29, 1898, Mill- spaugh 82 (F); Lake Superior, Parry (NY); Mackinac Island, 1888, Puckner (CAS); on Isle Royale and south shore of Lake Superior, Keweenah Point, 1862, Bobbins 102 166 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 20 (G, M); Keweenah Point, 1863, Robbins 95 (MU); without locality, Aug. 1887, Root (F); Chandler's Falls of the Escanaba River, Aug. 27, 1892, Wheeler (US); Thunder Bay Island, July 18, 1895, Wheeler (US); same locality and collector, Aug. 12, 1895 (NY); Harbor Springs, Aug. 10, 1890, Wheeler (F); Isle Royale, woods, mainland, Aug. 15-16, 1912, Williamson 2218 ( ANSP). *, sandy pine woods, Aug. 18, 1884, Hasse (ANSP, W I SCON bank 1893, Harper (M); Minah R., Door Co., July 27, 28, 29, 31, 1887, Schuette (F, G, K, NY, US); Europe Lake, Ellison Bay, Door Co., July 18, 1918, Stanton 19 (M); Cato Rapids, 1874, Swezey 17 (US). Illinois: Grand Detour, Porter (BB); same locality, Aug. 5, 1805, Smith (ANSP); same locality, Aug. 5, 18G6, Smith 5026 (BB). Minnesota: pine woods, Lake Itaska, July 1891, Aiton (BB, NY, US); Benedict, Norway-Jack pine forest, July 10, 1914, Bergman 2948 (NY); dry pine lands, Itaska Co., Aug. 1891, Burglehaus (M); Duluth, July 11, 1877, Hall (BB); Grand Marais, shore of the Bay and Lighthouse Point, Lake Superior, Aug. 9, 1920, Rydberg 9619 (NY); Lake Itaska, pine woods, July 1891, Sandberg (CAS, F); Two Harbors, July 1, 1891, Sandberg 457 (US); Grand Rapids, Aug. 6, 1891, Sandberg 719 (US); woods and shores, Lake Co., shady woods, Itaska Lake, Sandberg 1151 (F, US); Duluth, 1887, Yaxcy (US); north shore, Lake Superior, 1890, Wheeler A Jones 1054 (G); Fond du Lac, July 19, 1SS9, Woods (US). South Dakota: slate schist, under willows, North Rapid Ranger Station, alt. 1400 m., Black Hills National Forest, July 12, 1908, Murdoch 3072 (F, G, NY); woods south of Box Elder Creek, Lawrence Co., Aug. 3, 1924, Over 16145 (US); Custer, alt,. 1375 m., Aug. 15, 1892, Rydberg 878 (NY, US); Nashy, Black Hills, July 25, 1912, Visher 1557 (NY). Saskatchewan: 1857-8, Palliser's Brit. N. Am. Exped., Bourgeau (BG, G); Cumberland House Fort, Drummond (G, K). Montana: Columbia Falls, Mrs. J. J. Kennedy 38 (NY); same locality, July 17, 1892, Williams 903 (C, NY, US). Albekta: Rocky Mt. House, forest floor, open, Nordegg Distr., Sept. 24, 1928, Brinkman 3678 (NY). British Columbia: Kicking Horse Valley, near Field, alt. 1000 m., July 21, 1906, Brown 667 (ANSP. G. NY. US): Field. Julv 14. 1904. Farr (K): Rame loealitv. An*. Macoun 1904 Canada without Locality: 1838, Franklin cfc Douglas (DH); "foret prfis de Fort Elliee" (rare), Aug. 26, 1857, Bourgeau (K); " Terra Hudsonica," 1837, Grisebach (BG); June 1849, Leston (DH); MacNab (K); 1869, Macoun 77 (US); Percival (ANSP). Mexico: Ties Marias Mts., alt. 2375 m., Morelos, Dec. 16, 1907, Pringle 13971 (US); Rinc6ii, alt. 2300 m., Morelia, April 1909, Arshie 37 (US). The Rothrock specimen listed from Labrador is from an early collection made before the Canadian boundaries were perma- nently established. No definite locality is given, and it is probable that Labrador is incorrect and that the plant actually came from a point further south. This is the case of the specimen collected 1933] ALLEN — THE GENUS HALENIA 167 by Macfarlane and that by Martin as well, both, according to the label, from Caribou Island, Labrador. Caribou Island is now included in Nova Scotia. For this reason, it is doubtful if the area of distribution of the species proper extends as far north as Labrador. 23a. H. deflexa var. Brentoniana Gray, Syn. Fl. N. Am. ed. 2,2^ 127. 1886. Halenia Brentoniana Griseb. Gen. & Sp. Gent. 325. 1839; Dietrich, Syn. PI. 2: 918. 1840; Hook. Fl. Bor.-Am. 2: 67. pi. 156. 1840. Tetragonanthus deflexus var. Brentonianus Britt. in Mem. Torr. Bot. Club 5: 261. 1894; Britt. & Brown, 111. Fl. 3: 15. 1913. T. Brentonianus Heller, Cat. N. Am. PI. 6. 1898; Muhlen- bergia 1 : 2. 1900. Low plant, 0.3-1.5 dm. high; stem erect, much branched, nodes more approximate than in the species; leaves 3-5-nerved, the radical leaves similar to species, upper subsessile, oblong-lanceo- late; inflorescence a 3-flowered cyme with the center flowers on long pedicels; corolla purple, .8-1.0 cm. long, tube .3-5 cm. long; corolla-lobes ovate, acuminate, delicately veined, papillate; spurs broad or slender and subhorizontal ; calyx .5-.7 cm. long, segments elliptical, acuminate, 3-veined, papillate; stamens approximately .2 cm. long; anthers broadly ovate; filaments linear. Distribution: Labrador, southward to Nova Scotia and Quebec. Specimens examined: Labrador: Red Bay, Sept. 7, 1891, Bowdoin College 290 (G); on the gneiss plain, in sand, Blanc Sablon, Str. Belle Isle, July 30, 1910, Femald & Wiegand 3909 (G, K, NY); damp sand, Forteau, Belle Isle, July 30, 1910, Fernald, Wiegand & Kittredge S910 (G); 1842, Loving (G); Battle Harbor, July 5, 1926, SewaU & Weed (F); Aug. 10, 1895, Stearns (US); hills, Forteau, Aug. 8, 1893, Waghorne (G, M, US); Battle Harbor, Aug. 6, 1913, Williamson 559 (ANSP, NY); same locality, Williamson 547 (ANSP, NY); on dry sandy hillside, under 100 m., Cartwright, Sandwich Bay, July 31, 1926, Woodworth 357 (G). Newfoundland: "lieux humides ou sees et decouverts a pres du Pain de Su&ve, St. Pierre," Aug. 19, 1901, Louis-Arsene 365 (NY); "lieux humides ou sees, decouverts ou bois6s, mais plus gen6ralement sans les bois ou sous les buissons, St. Pierre, Pain de Sueve," Aug. 26, 1901, Louis-Arsene 403 (G); 1776, Banks (BM); without locality or date, Brenton 144a? (K); Bay St. George, dry sandy field along shore, Aug. 12, 1908, Eames & Godfrey 8030 (ANSP, G, K); turfy slopes of slaty hills, Little Quirpon, Quirpon Harbor, Aug. 6, 1925, Femald & Long 28950 (G, UC); boggy limestone barrens, Capstan Point, Flower Cove, Str. Belle Isle, July 28, 1924, Fernald, Long [Vol. 20 168 ANNALS OF THE MISSOURI BOTANICAL GARDEN & Dunbar 26982 (G); turfy limestone shore, sandy cove, Ingornachoix Bay, Aug. 9, 1924, Fcrnald, Long & Dunbar 269S3 (G, K); dry peaty barren, near Biscay Bay, Avalon Peninsula, Aug. 16, 1924, Fernald, Long A Dunbar 26984 (G); by rills on seepy silicioui slope of Joan Hill, Bay Bulls, Avalon Peninsula, Aug. 21, 1924, Fernald, Long A Dunbar 26985 (G); in turf on granite ledges, Gaultois southern coast, Aug. 29, 1924, Fernald, Long & Dunbar 26US6 (G); peaty and gravelly open slopes, French or Tweed Island, Bay of Islands, Sept. 2, 1926, Fcrnald, Long & Fogg 381 ((1); wet moss and peat on gneiss hills near sand bank west of Burges, Distr. of Burgos and LnPoile, Sept. 9, 1926, Fernald, Long & Fogg 382 (G); grassy fields over- lying conglomerate limestones and calcareous sandstones, Cowhead, Silurian coastal region north of St. Paul's Bay, July 22, 1910, Fcrnald gracilis 4. Steins 1-nmny; rosette present in complete plant. 6. Flowers 1-1.5 cm. long; plant yellow-green 43. U. Killipii 5. Flowers less than 1 cm. long (except in H. Weberbaueri); plant not yellow-green. G. Flowering stem scapiform, almost aphyllous; ilowers 5 or less. 44- H- Mathewsii 6. Flowering stem leafy; flowers more than 5. 7. Leaves linear, narrowly linear-lanceolate. 8. Plant less than 20 cm. high. 1933] ALLEN — THE GENUS HALENIA 181 9. Plant more than 10 cm. high; flowers .6-.7 cm. long. . . 45- H. vincetoxicoides 9. Plant less than 10 cm. high; flowers 1.2-1.5 cm. long 46. H. Weberbaueri 8. Plant more than 25 cm. high 47 H. Stuebelii 7. Leaves ovate-lanceolate to oblong-lanceolate; stem stout, rigid 48. H. robusta 2. Spurs conspicuous, }4~% the length of the corolla. 3. Stem decumbent, suffruticose; leaves subcoriaceous 49- H. taruga gasso 3. Stem usually erect, not suffruticose. 4. Flower 2-3 cm. long 50. H. gigantea 4. Flower less than 1.5 cm. long. 5. Leaves less than .8 cm. long 61. H. minima 5. Leaves more than .8 cm. long. 6. Flowering stems curved at apex 52. H. pendulifiora 6. Flowering stems erect. 7. Pedicels scarcely 1 cm. long; flowers in dense heads. 8. Cauline leaves oblong, hardly narrowed at base 53. H. phyteumoides 8. Cauline leaves lanceolate or oblong-lanceolate, gradually elongated into petiole, dilated at base 54. H. Herzogii 7. Pedicels more than 1 cm. long; flowers more loosely clustered. 8. Flowers less than 1 cm. long 55. H. silenoides 8. Flowers more than 1 cm. long. 9. Spurs slender, more or less parallel; plant more than 15 cm. high 56. H. umbellata 9. Spurs thick, distinctly incurved at tip; plant usually less than 15 cm. high 57. H. Meyeri Johannis 1. Spurs pendulous, divaricate. 2. Spurs scarcely % the length of the corolla, more or less divergent at tip. 3. Stems and branches densely and long-ciliate 58. H. barbicavlis 3. Stems and branches not ciliate, smooth, 4. Pedicels of apical flowers more than 1.5 cm. long. 5. Leaves ovate to obovate-lanceolate 59. H. Rusbyi 5. Leaves linear or linear-lanceolate. 6. Plant more than 10 cm. high, not caespitose 60. H. Purdieana 6. Plant less than 10 cm. high, densely caespitose 60a. H. Purdieana var. congesia 4. Pedicels of apical flowers less than 1.5 cm. long 61. H. Hieronymi 2. Spurs M-/4 the length of the corolla, horizontally or subhorizontally divaricate, but incurved at apex. 3. Stem strict; sterile branches none; basal leaves rigid and erect, dis- posed in a dense rosette 62. H. bifida 3. Stems more or less flexuous; sterile branches numerous; basal leaves not rigid, occasionally recurved, not disposed in a dense rosette . . . 63. H. Weddelliana 1. Spurs horizontal or reflexed. 2. Flowers 3-4 cm. long 64- H. elegans 2. Flowers less than 2 cm. long. 182 ANNALS OF THE MISSOURI HOTANICAL GARDEN [Vol. 20 3. Flowers less than I cm. long 65. II. Hoppii 3. Flowers more than 1 cm. long. 4. Sepals ovate-lanceolate. f>. Leaves thin, herbaceous, no rosette; spurs thick 66. H. asclepiadea 5. Leaves subcoriaceous; dense, many-leaved rosette; spurs slender. 67. II. Kalbreyeri 4. Sepals obovate or oblaneeolate. 5. Rosette and lower leaves lanceolate, slightly narrowed at base into a short petiole 68. II. bella 5. Rosette and lower leaves obovate, long-petiolate 69. II. sphagnicola 35. H. valerianoides CJilg in Engl. Bot. Jahrb. 54: Beibl. 118, p. 106. 1916. Si mill caespitose perennial, less than .5 dm. high; root very- thick and densely branching; stems short, erect, 2-3 cm. high, almost leafless, arising from a dense rosette of basal leaves; basal leaves thick, petiolate, oblaneeolate, 2-3 cm. long, .4 cm. broad, acute, obsoletely 3-nerved; cauline leaves broadly sessile, 1 pair or none, ovate-oblong, .5-.G cm. long, .2-.3 cm. broad; inflores- cence a terminal 1-3-flowered cyme, pedicels .7-1.0 cm. long; calyx-lobes obovate-oblong, .3 cm. long, .2 cm. broad, acute, obsoletely 3-nerved; corolla approximately .5 cm. long, .4 cm. broad, tube slightly less than one-half the length of the entire corolla; lobes ovate, acute; spurs small laterally prominent pro- tuberances at the base of the corolla. Distribution: Peru and Bolivia. Specimens examined: Peru: (type not seen, Weberbauer 1676, HG, M photo). According to Gilg, this number is incorrect, since the data given does not agree with the specimen. Bolivia: Chacaltaya, 130 klm. from La Paz, alt. 4800 m., Feb. 1908, Buchtien 1484 (US); Alaska Mine, alt. about 4500 m., March 1-4, 1926, Tate 67 (NY). 36. H. pusilla Gilg in Engl Bot. Jahrb. 54: Beibl. 118, p. 107. 1916. //. Dombeyana var. brevicornis Wedd. Chlor. And. 2: 76. 1859, fide Gilg, /. c. Perennial caespitose herb, under .5 dm. high; root short, thick, covered with remains of marcescent leaves, bearing at the apex a dense rosette of leaves; flowering stems numerous, densely crowded, erect, short; rosette leaves petiolate, very thick, ob- laneeolate, 1-1.5 cm. long, .2-.3 cm. broad, acute, obsoletely 3- nerved; cauline leaves 1-2 pairs, closely approximated, sessile, 1933] ALLEN — THE GENUS HALENIA 183 ovate-oblong to oblong, .6-7 cm. long, .3-35 cm. broad; inflores- cence a few-flowered (5 usually) cyme, pedicels up to 1 cm. long; calyx-lobes oblong-lanceolate, .5 cm. long, .2 cm. broad, acute, obsoletely 3-nerved; corolla .5-.6 cm. long, tube about one-half the length of the entire corolla; lobes ovate, acute; spurs small laterally prominent protuberances at the base of the corolla; stamens about .2 cm. long; filaments linear; anthers ovate, tip attenuate; capsule ovate. Distribution: Bolivia and Peru. Specimens examined: Bolivia: Prov. Larecaja, on road to Lacatia, in meadows, in vicinity of Sorata, 3200-3700 m.. Mandon MacBride 3600 (US) 37. H. spatulata 59 Allen, n. sp. Perennial caespitose herb, up to .8 dm. high; root coarse, woody; 1-2 flowering stems, erect, simple or rarely branched from base, occasional short sterile leafy branches ; numerous basal leaves attenuate into long petioles, elliptic to spatulate, up to 2.5 cm. long, .4-.5 cm. broad, prominently uninerviate; cauline leaves 1-2 pairs, sessile, elliptic, less than 1 cm. long; inflorescence usually terminal, 1 -few-flowered cyme, pedicels erect or slightly recurved at tip; calyx-lobes spatulate, up to .6 cm. long, .2 cm. broad, 3-nerved; corolla 1 cm. long, "lime green/' tube over one- half the length of the entire corolla; lobes broadly ovate, acute; spurs approximately one-fourth the length of the corolla, pendulous and incurved, broad at the base, attenuate at tip; stamens 69 H. spatulata Allen, sp. nov. — Herba perennis, caespitosa, usque ad .8 dm. alta; radice crassa, lignea; 1-2 caulibus floriferis, erectis, simplicibus vel raro e baso ramosis, vel ramis brevibus sterilibus foliosis; foliis basalibus multis, longis petiolis attenuatis, ellipticis vel spatulatis, usque ad 2.5 cm. longis, .4-.5 cm. latis, prominente 1-nerviis; foliis caulinis 1-2 geminis, sessilibus, ellipticis, minusquam 1 cm. longis; inflorescentia plerumque terminali, cymosa, 1-pauco-florifera; pedicellis erectis vel ad apicem parum recurvatis; calycis lobis spatulatis, usque ad .6 cm. longis, .2 cm. latis, 3-nerviis; corolla 1 cm. longa, "viride"; tubo plusquam M> corollae longitudini adaequanti; lobis late ovatis, acutis; calcaribus x /i corollae longitudini subaequanti, pendulis incurvatisque, ad basin latis, ad apicem attenuatis; staminibus ca. .2 cm. longis; filamentis linearibus; antheris ovatis; capsula late lanceolata; stigmatibus truncatis, planis superficiebus, ut videtur, stigmaticibus. — Peru: Dept. Cusco, open, grassy p&ramo, Cerro de Colquipata, alt. 3900-4000 m., May 1, 1925, Pennell 13749 (ANSP type, NY, US). [Vol. 20 184 ANNALS OF THE MISSOURI BOTANICAL GARDEN about .2 cm. long; filaments linear, anthers ovate; capsule broadly lanceolate; stigmas truncate, the flat surfaces apparently stig- matic. Distribution: Peru. Specimens examined: 3900-4000 13749 38. H. caespitosa Gilg in Fedde, Rep. Spec. Nov. 2: 53. 1906; ( Jilg in Engl. Bot. Jahrb. 54: Beibl. 118, p. 107. 1916. Small caespitose herb, forming a broad dense mat .5-8 dm. in diameter, usually .4-.5 dm. high, the flowering stems frequently reaching a height of 1-1.5 dm.; sterile branches usually intermin- gled with the fertile; leaves in dense aggregation, very fleshy, petiolate, obovate-lanceolate to oblanceolate, 1.5-2 cm. long, about .3-.4 cm. broad, acute, nerves scarcely conspicuous; cauline leaves, when present, sessile, oblong-lanceolate to linear- lanceolate, 1-2 cm. long; inflorescence usually consisting of a 1-, rarely 2-3-, flowered cyme at the apex of the fertile stem, pedicels 1.6-2 cm. long, erect or slightly nodding; calyx-lobes oblanceolate to oblong-lanceolate, .6-.8 cm. long, .2 cm. broad, acute to ob- tusish, 3-nerved; corolla about 1 cm. long, greenish, length of tube nearly equalling that of the entire corolla; lobes ovate, subrotund, crisped; spurs pendant, .1-2 cm. long and almost as thick; stamens approximately .4 cm. long, attached just below the sinus; filaments linear, anthers ovate; capsule linear. Distribution: moist places in Peru. Specimens examined: 3300-3G00 stream margin, Morococha, May 23, 1922, Macbrule A Featherstone 898 (F, M); "Hacienda Arapa bei Yauli, an der Lima-Oroya-Bahn," alt. 4400 m., 1906, Weber- bauer 27!) (BG type, DH, M photo). 39. H. hypericoides* (HBK.) G. Don, Gen. Hist. 4: 177. 1838; Griseb. Gen. & Sp. Gent. 328. 1839. Swertia hypericoides HBK. Nov. Gen. & Sp. PL 3: 176. 1818; Roem. & Schult. Syst. Veg. 6: 76. 1820. Perennial herb; stem procumbent, branching, leafy, less than 60 The specimen in the Herbarium of the Missouri Botanical Garden was collected by Margaret Kalenborn, No. 91. That from the United States National Herbarium bears the same number, but the collector is A. S. Kalenborn. In all probability these are the same collection. 1933] ALLEN — THE GENUS HALENIA 185 .3 dm. high; lower leaves several pairs, approximate, petiolate, sheathing at base, oblong-lanceolate or lanceolate, 2 cm. long, acute, 3-nerved; upper leaves smaller, sessile, oblong; inflorescence terminal and axillary many-flowered panicles, pedicels up to 1.5 cm. long; calyx-lobes linear-lanceolate, about .5 cm. long; corolla .7-.8 cm. long, yellowish; lobes ovate, acute; spurs pendulous, incurved, subconical, about one-half the length of the corolla; filaments linear, anthers oblong; capsule oblong, obtuse, com- pressed; seeds subglobose, blackish brown, smooth. Distribution: Colombia. No specimen examined, but description compiled from original publication and photograph, (type, Humboldt & Bonpland, HJP, M photo). 40. H. pulchella Gilg in Engl. Bot. Jahrb. 54: Beibl. 118, p. 107. 1916. Perennial herb, .5-1.0 dm. high; root thick, elongate; axis elongate, more or less procumbent, densely covered with obsolete remains of leaves, the apex curved-erect, sending out a single loosely leafy flowering stem, and some few (3-5) sterile pro- cumbent or curved-erect leafy branches 3-13 cm. long; rosette leaves lacking; lower cauline leaves broadly petiolate, upper sessile, rather fleshy, obovate, 1.2-1.6 cm. long, .5-.6 cm. broad, manifestly 3-nerved, veins sunken above, prominent below, acute; inflorescence composed of terminal and axillary pseudoracemose 3-flowered cymes, more or less approximate, pedicels .6-1.2 cm. long; leaves subtending inflorescence, minute, euphylloid; calyx- lobes oblong-lanceolate to oblanceolate, about .8 cm. long, .25- .3 cm. broad, 3-nerved, margin hirtellous; corolla about 1 cm. long, tube less than one-half the length of the entire corolla; lobes ovate, somewhat acute, auriculate; spurs pendulous, incurved, about one-half the length of the corolla; stamens about .2 cm. long; filaments linear, anthers ovate. Distribution: Ecuador. Specimens examined: Ecuador: in the Andes, Jameson 53 (DH type, M photo, V). Very similar to H. Weddelliana, but rather stouter, more leafy, and spurs less divergent. 41. H. pinifolia (R. & P.) G. Don, Gen. Hist. 4: 177. 1838. Swertia pinifolia (R. & P.) ex Don, I. c. 186 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 20 Perennial herb, growing in tufts, .7-1.5 cm. high; steins erect, simple; leaves linear, channeled, acute, edges scabrous; inflores- cence umbellate; calyx-lobes ovate-lanceolate, acute; corolla golden-yellow; spurs straight, one-half as long as the corolla. Distribution: Cordilleras of the Andes of Peru. No specimens examined, but description compiled from original publication. (type, Ruiz & ravon — Herbarium at Madrid). 42. H. gracilis (HBK.) G. Don, Gen. Hist. 4: 177. 1838; Griseb. Gen. & Sp. Gent. 327. 1839; DC. Prodr. 9: 130. 1845 (exel. var.). Swertia gracilis HBK. Nov. Gen. & Sp. PL 3: 170. 1818. Halcnia pichinchensis Gilg in Engl. Bot. Jahrb. 54: Beibl. 118, p. 109. 1916. H. Jamesoni Gilg in Engl. Bot. Jahrb. 54: Beibl. 118, p. 115. 1916. Annual herb, up to 3 dm. high; stem simple, erect, minutely striate; few basal leaves suggesting a rosette, usually fugacious, with persistent bases, broadly elliptic to ovate, attenuate into narrow petioles longer than the blade, up to 3 cm. long, up to 1 cm. broad, acute, prominently 3-nerved; cauline leaves up to 5 pairs, at intervals of 4 cm., petiolate, elliptic, becoming broadly lanceolate toward the summit of the stem, acute, 3-nerved; floral leaves smaller than cauline; inflorescence few-flowered (2- 6) axillary and terminal cymes, erect pedicels up to 1.2 cm. long; calyx-lobes more or less obovate, papillate, approximately .6 cm. long, .2 cm. broad, abruptly acuminate, 3-nerved, reticulate at tip; corolla 1-1.5 cm. long, greenish, tube one-half the length of the entire corolla; lobes ovate, acuminate; spurs slightly less than one-third the length of the corolla, very slender, slightly diver- gent, the tip frequently, but not always, slightly incurved; stamens approximately .3 cm. long, attached just below the summit of the tube; filaments linear, anthers oval, abruptly acuminate; stigmas broad, recurved; capsule lanceolate, sub- falcate, 1.8 cm. long; seeds reticulate. Distribution: Colombia and Ecuador. Specimens examined: Colomhia: Zipaquira, alt. 2730 m., Humboldt & Bonpland (HJP type, M photo). Ecuador: Pichincha, Karstcn (BG, V); on Mt. Pichincha, opposite Quito, March 3, 1920, Ilcilborn 437 (V); La Planta del Chillo, about Tanque, alt. 2700 m., April 1933] ALLEN — THE GENUS HALENIA 187 2, 1920, Firmin 697 (US); near Quito, coll. of 1864, Jameson (V); about Quito, Jameson (ANSP, BG); Guagrapata, Spruce 5131 (V). Halenia gracilis shows a marked relationship to Halenia Schiedeana of Mexico. The specimens which Gilg has described as H. Jamesoni and H. pichinchensis appear to be conspecific with gracilis. In the opinion of the author, the only possible difference is the texture of the leaves, which in the Jamesoni specimen is slightly rougher than that of the gracilis type. 43. H. Killipii 61 Allen, n. sp. Pale green perennial, .3-2.5 dm. high; root coarse, heavy, ligneous, covered with darkened remains of leaves; stem stout, conspicuously alate; basal leaves few, fleshy, in rosette, attenuate into long petioles, oblanceolate, 2-4 cm. long, .3-35 cm. broad, 3-nerved, obtuse; cauline leaves 1-3 pairs, sessile, linear-lance- olate, 1.5-2.5 cm. long, .3 cm. broad, inconspicuously 3-nerved, obtuse; inflorescence 1 (rarely 7)-flowered, usually terminal, subumbellate cyme, pedicels erect or slightly nodding, up to 2.5 cm. long; calyx-lobes elliptic, .5-.9 cm. long, yellowish-green, acute, obsoletely 3-nerved; corolla 1-1.5 cm. long, tube about one- third the length of the entire corolla; lobes broadly ovate, erose; spurs thick, pendulous, incurved, about one-third the length of the corolla; stamens about .2 cm. long; filaments linear, anthers ovate; capsule ovate, attenuate at apex, yellowish-green. Distribution: Peru. Specimens examined: Peru: Dept. Junfn, Mount La Juntay, near Huancayo, alt. 4700 m., April 27, 1929, Killip & Smith 22087 (US type); same locality and date, Killip & Smith 22083 (US). 61 H. Killipii Allen, sp. nov. — Herba perennis, pallida viride, .3-2.5 dm. alta; radice crassa, lignea, reliquiis foliorum tecta; caule robusto, conspicue alato; foliis basalibus paucis, carnosis, in rosula, petiolis longis attenuatis, oblanceolatis, 2-4 cm. longis, .3-35 cm. latis, 3-nerviis, obtusis; foliis caulinis 1-3 geminis, sessilibus, lineari-lanceolatis, 1.5-2.5 cm. longis, .3 cm. latis, inconspicue 3-nerviis, obtusis; inflorescentia 1 (raro 7)-florifera, plerumque terminali, subumbellata-cymosa; pedicellis erectis vel parum nutantibus, usque ad 2.5 cm. longis; calycis lobis ellipticis, .5-.9 cm. longis, flavo-viridibus, acutis, obsolete 3-nerviis; corolla 1-1.5 cm. longa; tubo ca. Yz corollae longitudini adaequanti; lobis late ovatis, erosis; calcaribus crassis, pendulis, incurvatis, ca. }/& corollae longitudini adaequanti; staminibus .2 cm. longis; filamentis linearibus, antheris ovatis; capsula ovata, apice attenuati, flavo- viride. — Peru: Dept. Junin, Mt. La Juntay, near Huancayo, alt. 4700 m., April 27, 1929, Killip & Smith 22087 (US type); same locality and date, Killip & Smith 22083 (US). 188 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 20 44. H. Mathewsii Gilg in Engl Bot. Jahrb. 54: Beibl. 118, p. 111. 191(>. H. asclepiadea Griseb. Gen. & Sp. Gent. 320. 1839; in DC. Prodr. 9: 129. 1845, pp; non Svrrtia asclepiadea HBK. fide Gilg, I. c. Perennial herb, up to 2.5 dm. high (usually less than 1.5 dm.); subterranean axis thick, short, erect; numerous flowering stems erect, slender, sparingly leafy; basal leaves in dense rosette, herbaceous, petiolate, obovate-oblong, up to 2.5 cm. long, .4-.5 cm. broad, the lowest equal and equidistant on the stem, the uppermost sessile and much reduced, oblong or ovate-oblong, more or less acute, obsoletely or inconspicuously 5-nerved; in- florescence a 5-flowered cyme, on a more or less scapiform stem; pedicels of terminal flowers 2 cm. long, of laterals 1.5 cm., de- creasing toward the base; calyx-lobes oblanceolate, about .55 cm. long, acute, obsoletely 3-nerved; corolla about .7 cm. long, tube about one-third the length of the entire corolla; lobes ovate- oblong, subrotund; spurs pendulous, slightly incurved, one-third the length of the corolla. Distribution: Peru. Specimens examined: Pkhu: near Huamatanga, Mathews 523 (V type); Dept. Lima, swale on p&ramo, near Antaicocha, Cerro Colorado, east of Canta, alt. 4000-4200 m., June 20, 1925, Pennell I ',67H (ANSP, NY, S, US); Dept. Lima, open hillside, Rio Blanco, alt. 3000-3500 m., April 15-17, 1929, Killip & Smith 21737 (US). 45. H. vincetoxicoides Gilg in Engl. Bot. Jahrb. 54: Beibl. 118, p. 108. 1916. Tetragonanthus Whitei Rusby in Mem. N. Y. Bot. Gard. 7: 321. 1927. Perennial herb, 1.5-2 dm. high; stems 1-5, simple, arising from tuft of basal leaves which may or may not be persistent, internodes up to 8 cm. long; basal leaves numerous, attenuate into long petioles, narrowly elliptic, 1.5-2.5 cm. long, .2-.45 cm. broad, acute; cauline leaves 2-3 pairs, sessile, subconnate, lance- olate, 1.5-3.5 cm. long, up to .4 cm. broad, 3-nerved, midvein prominent; inflorescence axillary and terminal many-flowered (6- 14) subumbellate cymes; pedicels usually recurved at tip, 2.5 cm. long; calyx-lobes oblanceolate-elliptic, papillate, up to .55 cm. long, .25 cm. broad, acute, 3-nerved, reticulate at tip; corolla 1933] ALLEN — THE GENUS HALENIA 189 .6-7 cm. long, yellowish, tube one-half the length of the entire corolla; lobes ovate-oblong, papillate, acute, margin inrolled and erose; spurs one-fourth length of corolla, slender, slightly spreading, incurved at tip; stamens approximately .3 cm. long, attached at summit of tube; filaments linear, anthers broadly ovate; stigma only slightly reflexed, capsule up to 1 cm. long, subf alcate ; seeds subglobose, reticulate, greenish yellow, brown. Distribution: moist grassy meadows of Bolivia. Specimens examined: Bolivia: Yungas, 1890, Bang 665 (BG type, G, M, NY, US, V); Sorata, alt. 3300 m., Feb. 1886, Rusby 669 (G, M, NY, US, V); Pongo, alt. 3800 m., July 11, 1921, White 178 (NY type of Tetragonanthus Whitei Rusby). This last-cited specimen was described as Tetragonanthus Whitei Rusby, but accords exactly with H. vincetoxicoides Gilg, excent for its small size. 46. H. Weberbaueri 62 Allen, n. sp. Perennial herb, caespitose, .5-1 dm. high; root fibrous, covered ith darkened remains of leaves; stems usually several, rather out : basal leaves numerous, attenuate into long slender petioles cm the blade in length, oblanceolate to lanceolate, £ .2-.4 cm. broad, obtuse, 3-nerved; cauline leaves s extreme base of stem, linear-lanceolate, 3^1 cm .2-3 cm. broad, obtuse, 3-nerved; inflorescence 3-5-flowered terminal subumbellate cymes with occasional depauperate 1- flowered cymes in axils of cauline leaves, pedicels .6-2 cm. long, the tips; calyx-lobes broadly oblanceolate 62 H. Weberbaueri 5-1 radice fibrata, reliquiis foliorum tecta; caulibus plerumque pluribus, aliquid robustis; foliis basalibus multis, petiolis longis tenuibus laminae longitudini adaequantibus 5-3 3-4 .2-3 cm. latis, obtusis, 3-nerviis; inflorescentia 3-5 florifera, terminali, subumbellata- cymosa, per occasionem depauperata 1-florifera foliorum caulinium axillibus; pedi- urvatis ■nervus 1.2-1.5 cm. longa, viridula; tubo circiter X A totae corollae longitudini adaequanti; lobis ovatis, acutis, erosis; calcaribus }4 corollae longitudini adaequantibus, tenuibus, pendulis, saepe parum divaricatis, sed semper ad apicem incurvatis ; staminibus minus- quam .2 cm. longis; filamentis linearibus, antheris ovatis; capsula lanceolata, ad apicem attenuata.— Peru: rocks, Mt. Razuhuillca, Pro v. Huanta, Dept. Ayacucho, alt. 4300-4500 m., Feb. 4-6, 1926, Weberbauer 7A98 (F type). [Vol. 20 190 ANNALS OP THE MISSOURI BOTANICAL GARDEN narrowly obovate-elliptic, .4-.6 cm. long, acute to obtuse, 3- nerved; corolla 1.2-1.5 cm. long, greenish, tube about one-half the length of the entire corolla; lobes ovate, acute, erose; spurs one-third the length of the corolla, slender, pendulous, often slightly divaricate but always incurved at the tips; stamens less than .2 cm. long; filaments linear, anthers ovate; capsule lanceo- late, attenuate at tip. Distribution: Peru. Specimens examined: Peru: rocks, Mt. Razuhuillca, Prov. Huanta, Dept. Ayaeucho, alt. 4300-4500 m., Feb. 4-6, 1926, Weberbauer 7498 (F type). 47. H. Stuebelii Gilg in Engl. Hot. Jahrb. 54: Beibl. 118, p. 110. 1916. Biennial, up to 3-4 dm. high; stem thick, erect, simple or branching at base; basal leaves in dense rosette, narrowed at base, lanceolate, 3-3.5 cm. long, .3-4 cm. broad, acute; cauline leaves thickly herbaceous, sessile, oblong-lanceolate, 2-5 cm. long, .3 cm. broad, the uppermost smaller, obsolete, narrowly acuminate, 5-nerved, veins sunken above, prominent below; inflorescence a terminal dense many-flowered (5-7-9) subcapitate cyme, and axillary 1 -few-flowered (3, rarely 5) cymes, pedicels 1-2.7 cm. long; calyx-lobes lanceolate, .7-8 cm. long, .25 cm. broad, acute, obsoletely 3-5-nerved; corolla almost. 1 cm. long, tube approxi- mately one-third the length of the entire corolla; lobes ovate- oblong, acute; spurs pendulous, incurved, about one-third the length of the corolla. Distribution: Peru. Specimens examined: Peru: Rio Blanco, about 5000 m., May 20-25, 1923, Macbrule 3040 (F, M). (type not seen, Sttibel Jfie, BG, M pboto). 48. H. robusta Gilg in Engl. Bot. Jahrb. 54: Beibl. 118, p. 110. 1916. Biennial herb, up to 3 dm. high; ligneous unbranched tap-root; stems 1-3, simple or branching below or above, erect, sturdy, up to .3 cm. thick, striate, internodes up to cm. long; numerous basal leaves in rosette, attenuate into narrower petiole, broadly elliptic, 2-2.5 cm. long, .6 cm. broad, acute-acuminate. 3-nerved. 1933] ALLEN — THE GENUS HALENIA 191 appearing sunken from above, prominent below; cauline leaves 1 cm. lone, .65 cm. broad 3-5-nerved, acute; inflorescence axillary and terminal many- flowered (6-15) subumbelliform cymes; pedicel sturdy, slightly recurved at tip, up to 3 cm. long; calyx-lobes oblong to ovate, approximately .5 cm. long, .25 cm. broad, acute, papillate, uninerviate, tip reticulate; corolla up to .7 cm. long, yellow, tube slightly less than one-half the length of the entire corolla; lobes narrowly ovate, acute ; spurs borne at the midpoint of the corolla- tube, tiny, slender, scarcely one-fourth the length of the corolla, tip incurved; stamens about .25 cm. long, inserted at the summit of the tube; filaments linear, anthers ovate-oval; capsule up to 1.3 cm. long, ovate; seeds oval, reticulate. Distribution: near snow line in Bolivia. Specimens examined: Bolivia: near snow line, Mt. Tunari, near Cochabamba, 1891, Bang 1019 (ANSP, BG type, F, G, M, NY, US, V); Dept. Cochabamba, Prov. Chapar6, Ceja-region, La Aduana, alt. 3000 m., March 7, 1927, Steinbach 9535 (M, NY, S). 49. H. taruga gasso Gilg in Engl. Bot. Jahrb. 54: Beibl. 118, p. 117. 1916. Suffruticose perennial, ligneous, more or less decumbent, up to 1.5 dm. high; root frequently sending up one or more short sterile densely leafy stems; flowering stems simple, leafy, rather slender; basal leaves, if present, petiolate, subcoriaceous, in dense spirals about lower portion of stem, lanceolate or linear- lanceolate, up to 2 cm. long, .35 cm. broad, acute; cauline leaves more or less sessile, more linear, otherwise similar to basal leaves ; inflorescence usually a terminal many-flowered (6) cyme, pedicels up to 1 cm. long; calyx-lobes oblong-elliptic, acute, 3-nerved, reticulate at tip; corolla up to 1.3 cm. long, greenish-yellow, tube approximately one-fourth the length of the entire corolla; lobes ovate, acute; spurs almost one-half the length of the corolla, slender, spreading, tip incurved or divergent; stamens about 2 cm. long; filaments linear, anthers ovate; capsule oblong-ovate, subfalcate, up to 1.3 cm. long; seeds subglobose, reticulate. Distribution: Ecuador. Specimens examined: Ecuador: Loja, alt. 3000-3500 m., Lehmann 4878 (BG type, M photo); Canar, Sept. 15, 1918, Rose & Rose 22674 (NY, US); Mt. Pittshum, Jameson (G). 192 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 20 50. H. gigantea r>3 Allen, n. sp. Stout coarse perennial, up to 4 dm. high; root thick, ligneous, up to .5 cm. in diameter; axis subhorizontal or horizontal, coarse, fleshy, .5-.(> cm. in diameter, covered with dense coarse black remains of leaves; stems 1-many, coarse, erect, simple, inter- nodes up to 7 cm. long; numerous basal leaves, in dense rosette, herbaceous, narrowed into petioles equalling the blade in length, di- lated ami almost sheathing at the base, lanceolate, 5-10 cm. long, .6-1 cm. broad, attenuate-acuminate, 3-5-nerved; cauline leaves about ii pairs, sessile, lanceolate, 2-3 cm. long, decreasing in length toward the summit, .5 cm. broad, acuminate, 3-nerved; inflorescence a loose terminal usually 3-flowered cyme, pedicels up to 5 cm. long, erect; calyx-lobes ovate to ovate-oblong to oblong, up to 1.5 cm. long, .5 cm. broad, acute, often with sug- gestion of apicule, 3-many-nerved ; corolla 3^1 cm. long, light green, tube less than one-third the length of the entire corolla; lobes oblong-ovate, acute, more or less erose, frequently sub- apiculate; spurs coarse, about one-third the length of the corolla, pendulous, incurved, the tip heavily glandular; stamens nearly 1 cm. long; filaments linear, anthers ovate, acuminate; capsule lanceolate. Distribution: Colombia. Specimens examined: Colombia: Dept. Santander, Pdramo de Santurbdn, near Vetas, alt. 3950-4160 H. gigantea Allen, sp. nov. — Herba perennis, robusta, crassa, usque ad 4 dm. alta; radiee crassa, lignea, usque ad .5 cm. in diametro; axe subhorizontale horizont- aleve, crasso, carnoso, usque ad .5-6 cm. in diametro, reliquiis foliorum densis crassis tec to; caulibus 1-multis, crassis, erectis, simplieibus, internodiis usque ad 7 cm. longis; foliis basalibus multis, in rosula densa, herbescentibus, petiolis laminae longitudini adaequantibus attenuatis, dilntatis et prope ad basin vaginantibus, lanceolatis, 5-10 cm. longis, .6-1 em. latis, attenuate acuminatis, 3-5-nerviis; foliis caulinis ca. 3 geminis, sessilibus, lanceolatis, 2-3 cm. longis, decrescentibus sursum, .5 cm. latis, acuminatis, 3-nerviis; inflorescentia laxa, cymosa, terminali, plerumque 3-florifera, pedicellis usque ad 5 cm. longis, erectis; calycis lobis ovatis vel ovato- oblongis vel oblongis, usque ad 1.5 cm. longis, .5 cm. latis, acutis, saepe subapicu- latis, 3-nmlti-nerviis; corolla 3-4 cm. longa, pallida viride; tubo minusquam \i corollae longitudini adaequanti; lobis oblongo-ovatis, acutis, plus minusve erosis, saepe subapiculatis; calcaribus crassis, ca. l /i corollae longitudini adaequantibus, pendulosis, incurvatis, ad apicem dense glandulosis; staminibus ca. 1 cm. longis; filamentis linearibus; antheris ovatis, acuminatis; capsula lanceolata. — Colombia: Dept. Santander, P&ramo de Santurb;in, near Vetas, alt. 3950-4160 m., Jan. 17 1927, Killip & Smith 17566 (M typb:, NY, US). 1933] ALLEN — THE GENUS HALENIA 193 m., Jan. 17, 1927, Kiilip & Smith 17566 (M type, NY, US); same locality, Killip & Smith 17521 (US); same locality, Killip & Smith 17516 (M, NY, US). 51. H. minima 64 Allen, n. sp. Small perennial, caespitose, .6 dm. or less high; root ligneous; frequently numerous short sterile leafy stems up to 2.5 cm. high; flowering stems 1-2 cm. high, erect, slender, almost scapiform, simple; basal leaves in dense rosette, thick, coarse, attenuate into petioles shorter than blades, broadly oblanceolate, less than 1 cm. long, .2-3 cm. broad, acute, obsoletely 3-nerved; leaves of sterile branches abruptly narrowed into petioles, exceeding blades in length, elliptic, up to .8 cm. long, .2 cm. broad, acute, obsoletely 3-nerved, midvein prominent; cauline leaves 1 pair at base, a second pair subtending the inflorescence, sessile, oblanceolate, .3-.7 cm. long, .2 cm. broad, acute to acuminate, obsoletely nerved; inflorescence a 2-4-flowered terminal loose cyme, pedicels 1.5 cm. long, slightly curved at apex; calyx-lobes oblong- oblanceolate, .4-. 5 cm. long, .1-.15 cm. broad, acute, obsoletely 3-nerved; corolla .6-.8 cm. long, tube one-half the length of the entire corolla; lobes ovate, acute; spurs slender, pendulous, slightly incurved, between one-third and one-half the length of the corolla; stamens about .2 cm. long; filaments linear, anthers ovate; capsule ovate, attenuate at apex. Distribution: Ecuador. Specimens examined: Ecuador: Andes, coll. of 1855, Couthouy (G type, NY). 64 H. minima Allen, sp. nov. — Herba perennis, parva, caespitosa, .6 dm. minusve alta; radice lignea; saepe caulibus multis, brevibus, sterilibus, foliosis, usque ad 2.5 cm. altis; caulibus floriferis 1-2 cm. altis, erectis, tenuibus, prope scapiformibus, simplicibus; foliis basalibus in rosula densa, crassis, petiolis brevioribus laminis attenuatis, late oblanceolatis, minusquam 1 cm. longis, .2-.3 cm. latis, acutis, ob- solete 3-nerviis; foliis ramorum sterilium petiolis longioribus laminis subito attenuatis, ellipticis, usque ad .8 cm. longis, .2 cm. latis, acutis, obsolete 3-nerviis, medio-nervo prominenti; foliis caulinis ad basin 1-geminis, alio gemine inflorescentiam subtendenti, sessilibus, oblanceolatis, .3-7 cm. longis, .2 cm. latis, acutis vel acuminatis, obsolete 3-nerviis; inflorescentia 2-4-florifera, terminali, laxa, cymosa, pedicellis 1.5 cm. longis, ad apicem parum curvatis; calycis lobis oblongo-oblanceolatis, .4-.5 cm. longis, .1-15 cm. latis, acutis, obsolete 3-nerviis; corolla .6-.8 cm. longa; tubo J^ corollae longitudini adaequanti; lobis ovatis, acutis; calcaribus tenuibus, pendulis, parum incurvatis, H~"3^ corollae longitudini adaequantibus; staminibus ca. .2 cm. longis; filamentis linearibus, antheris ovatis; capsula ovata, ad apicem attenuata. — Ecuador: Andes, coll. of 1855, Couthouy (G type). 194 ANNALS OF THE MISSOUEU HOTANICAL GARDEN [Vol. 20 52. H. penduliflora (iilg in Engl. Bot. Jahrb. 54: Beibl. 118, p. 110. 1916. H. Dombeyana var. a Wedd. Chlor. And. 2 : 7(5. 1859 pp. Biennial, up to 3 dm. high; root fibrous, axis thick, short, covered with remains of marcescent leaves; stem single, simple, erect, curved at apex; basal leaves in loose rosette, herbaceous, petiolafe, oblong-lanceolate to elliptic, 4 cm. long, ,5-.6 cm. broad, acute; cauline leaves 3 pairs or so, slightly narrowed at base, but sessile, oblong, 2-3 cm. long, .5-.7 cm. broad, acute, 3- nerved ; inflorescence a 7-11-ilowered subumbellate terminal cyme, always pendulous, rarely an axillary small 3-flowered cyme, pedicels 1.5 cm. long ; calyx-lobes oblanceolate to oblong-oblanceo- late, about .8 cm. long, .2-5 cm. broad, obsoletely 3-nerved; corolla 1-1.2 cm. long, tube scarcely one-third the length of the entire corolla; lobes ovate, acute; spurs pendulous, slightly di- vergent, but incurved at tips about one-third to three-fourths the length of t he corolla; stamens about .4 cm. long; filaments linear, anthers ovate; capsule ovate, attenuate at apex. Distribution: Bolivia. Specimens examined: 3200-3700 M photo, NY, 8, V). The specimen of Halenia pendulijlora Gilg, based on Mandon 869 pp., has the same floral characteristics as silenoides and bears the label "in graminosis," which signifies a possible ecological variation. The habit is that of typical specimens of silenoides, which, having grown in grassy situation, has become attenuate, with the internodes more elongate, the stem more or less decum- bent. However, until more material is available the species pendulijlora must be retained. 53. H. phyteumoides Gilg in Engl. Bot. Jahrb. 54: Beibl. 118, p. 112. 1916. Perennial caespitose herb, up to 1.2 dm. high; root short, thick, densely fibrous, with very short axis; stem lateral, erect, simple, thick, angled; basal leaves in dense rosette, attenuate into long petioles equalling the blades in length, spatulate or broadly oblanceolate, up to 2.5 cm. long, .35 cm. broad, acute, obsoletely 3-nerved; cauline leaves 1-2 pairs, slightly narrowed at base, but 1933] ALLEN — THE GENUS HALENIA 195 sessile, thick, 1-1.5 cm. long, decreasing toward the summit, .3- .4 cm. broad, acute, obsoletely 3-nerved; inflorescence a terminal 5-flowered subcapitate cyme and axillary 3-flowered cymes; pedicels strongly winged, the apical up to 1 cm. long, the laterals .5-.6 cm. long; calyx-lobes obovate-oblong, up to .4 cm. long, .2 cm. broad, acute, obsoletely 3-nerved; corolla .8 cm. long, tube slightly less than one-half the length of the entire corolla; lobes ovate-oblong, subrotund; spurs slender, pendulous, incurved, one-half the length of the corolla. Distribution: Peru. No specimens examined, but description compiled from original publication and photographs, (type, Philipj/i, BG, M photo). 54. H. Herzogii Gilg in Engl. Bot. Jahrb. 54: Beibl. 118, p. 113. 1916. Perennial caespitose herb, up to .5 dm. high ; subterranean axis thick, black, branching, elongate, covered with blackened re- mains of leaf bases ; stem curved-erect, scapiform, strongly winged, simple; basal leaves in dense rosette, thick, fleshy, attenuate into long petioles equalling the blades, spatulate or obovate-oblong to oblanceolate, up to 2.6 cm. long, .3 cm. broad, acute or more or less rounded at apex, obsoletely 3-nerved; cauline leaves 1-2 pairs, attenuate, but dilated at base and sessile, lanceolate to oblong-lanceolate, 1-2 cm. long, decreasing toward the summit, .25-.3 cm. broad, acute, obsoletely 3-nerved; inflorescence a small terminal 5-flowered subcapitate cyme; terminal pedicels 1 cm. long, lateral .5-. 6 cm. long, strongly winged; calyx-lobes obovate- oblong to broadly oblanceolate, about .4 cm. long, less than .2 cm. broad, acute, obsoletely 3-nerved; corolla .6-7 cm. long, tube about one-half the length of the entire corolla; lobes narrowly ovate, subrotund; spurs one-third to three-fourths the length of the corolla, pendulous, incurved; stamens about .15 cm. long, inserted at the orifice of the tube; filaments linear, anthers ovate; stigma truncate; capsule lanceolate. Distribution: Bolivia. Specimens examined: Bolivia: Lagodos, alt. 4400 m., Herzog 2S77 (BG type, M photo, V). 55. H. silenoides Gilg in Engl. Bot. Jahrb. 54: Beibl. 118, p. 113. 1916. [Vol. 20 190 ANNALS OF THE MISSOURI BOTANICAL GARDEN Halenia Dombeyana var. a Wedd., Chlor. And. 2 : 76. 1859, pp. non Halenia gracilis var. Dombeyana Griseb. Perennial herb, for the most part less than 1.5 dm. high, seldom over 2 dm.; root thick, many branches; stems 1-6, simple, erect or very slightly decumbent, rather thick, covered with marcescent leaves at the bases; sterile stems short, densely leafy; basal leaves numerous, attenuate into long narrow petioles frequently equalling the blades, less than 2 cm. long, .35 cm. broad, acute, 3-nerved; cauline leaves 1-2 pairs, sessile, lanceolate, usually about 2 cm. long, acute, 3-nerved; inflorescence a terminal or axillary 3-10-flowered cyme, pedicels more or less erect, up to 2.5 cm. long; calyx-lobes oblong-oblanceolate, approximately .3-.5 cm. long, acute, uninerviate, densely reticulate; corolla approximately .9 cm. long, yellowish-green, tube about one-half the length of the entire corolla; spurs more than one-third the length of the corolla, slender, divergent, incurved at tips ; stamens about .25 cm. long; filaments linear, anthers broadly ovate, acute; capsule up to 1.1 cm. long, narrowly ovate, attenuate at tip, subfalcate; seeds subglobose, reticulate. Distribution: alpine meadows in Bolivia and Peru. Specimens examined: Bolivia: alpine meadows, Choquetanga Grande, alt. 3600 m., Herzog 2402 (S); Unduavi, Nordyungas, 3300 m., Buchlien 54 (F, G, NY); Unduavi, alt. 3300 m., Buchtien 600 (US) ; Unduavi Valley, Bro. Jvlio 838 (US) ; same locality, alt. 2000- 2600 m., 1925, Bro. Jvlio 455 (US); Pongo, alt. 4000 m., Feb. 17-March 1, 1926, Tate 223 (NY); Mandon 369 pp (BG type?). Bolivia without Locality: Cumming 128 (V). Peru: Dept. of Cusco, Paso de Tres Cruces, Cerro de Cusilluyoc, alt. 3800-3900 m., May 3, 1925, Pennell 13842 pp. (ANSP, US). The specimen collected by Tate is rather doubtfully included. It has the habit of silenoides, but the flowers on the whole appear smaller, the spurs shorter and thicker, more like those of Her- zogii. The specimens, Pennell 188 J$, found in the United States National Herbarium and the Philadelphia Academy of Natural Sciences, are certainly H . silenoides, but the same number located in the New York Botanical Garden Herbarium is H. asclepiadea. 56. H. umbellata (R. & P.) Gilg in Fedde, Rep. Spec. Nov 2:53. 1906. Swertia umbellata R. & P. Fl. Peruv. 3: 21. pi. 21$, fig. b 1802. 1933] ALLEN — THE GENUS HALENIA 197 Halenia Pavoniana G. Don, Gen. Hist. 4: 177. 1838. Halenia gracilis var. /3 Dombeyana Griseb. in DC. Prodr. 9: 130. 1845. Halenia Dombeyana Wedd. Chlor. And. 2: 76. 1859. Perennial herb up to 3 dm. high; root ligneous, frequently- sending out short sterile densely leafy branches; stems 1-3, usually simple, erect, minutely striate, frequently branched above, the branches bearing inflorescences nearly as long as the terminal branch, marcescent leaves at base; basal leaves numer- ous, elliptic, attenuate into slender petioles nearly equalling blades, 2-3.5 cm. long, .6-.8 cm. broad, acute, 3-nerved; cauline leaves 2-3 pairs, attenuate into short petioles, or the extreme upper more or less sessile, lanceolate to elliptic, acute, 3-nerved; inflorescence terminal or axillary 5-14-flowered umbellate cymes, with an approach to a corymb; pedicels up to 3.5 cm. long, the center usually erect, the marginal more or less recurved and shorter; calyx-lobes obovate-elliptic, up to .6 cm. long, sub- acuminate, 3-nerved, reticulate at tip; corolla 1.0-1.3 cm. long, tube slightly less than half the length of the entire corolla; lobes ovate-oblong, acutish; spurs less than one-half the length of the corolla, very slender, tapering, slightly divergent; stamens .2-25 cm. long; filaments linear, anthers broad-oblong, somewhat acute; stigma deeply cleft, slender, attenuate; capsule up to 1.5 cm. long, narrowly ovate-attenuate, subfalcate; seeds globose, reticulate. Distribution: Andes of Peru. Specimens examined: Peru: Baflos, Dombey (G, US); Agapata in Virgallis, coll. of 1854, Lechler 2001 (DH, V); Lucumayo Valley, alt. 1800-3600 m., June 18, 1915, Cook & Gilbert 1S08 (US); Pifiasniocj, Panticalla Pass, alt. 3600 m., July 14, 1915, Cook & Gilbert 1811 (US); Mito, alt. 3000 m., July 8-22, 1922, Macbride & Featherstone 1657 (F, M); La Quinua, alt. about 4000 m., May 14, 1922, Macbride & Featherstone 2001 (F, M); DeDt. of Cusco. Cerro de Colauioata. alt. 4100-4200 m.. May 1. 1925, Pennell 1S738 2700 m.. Weberb UT The illustration of S. umbellata in Ruiz & Pavon's 'Flora jruvianae' has spurs longer than the specimens cited above but e nlant is similar otherwise. 57. H. Meyeri Johannis Gilg in Engl. Bot. Jahrb. 54: Beibl. 118, p. 114. 1916. [Vol. 20 198 ANNALS OF THE MISSOUIU BOTANICAL GARDEN Perennial herb up to 2 dm. high, though usually less than 1.5 dm.; root ligneous; stem simple, erect, rather stout base frequently covered with marcescent leaves; basal leaves, when present, in dense rosette, attenuate into petiole longer than blade, narrowly elliptic, up to 2.2 cm. long, up to .6 cm. broad, acute, 3-nerved; 1-2 pairs of cauline leaves, sessile, elliptic, up to 1.8 cm. long, about .5 cm. broad, inconspicuously 3-nerved, acute; inflorescence an axillary or terminal, 3-12-flowered cyme, loose or dense; pedicels up to 3 cm. long, usually recurved; calyx-lobes obovate-elliptic, papillate, approximately .(i cm. long, .2 cm. broad, 3-nerved, acute, reticulate; corolla up to 1.5 cm. long, yellowish, tube less than one-half the length of the entire corolla; lobes ovate-acute; spurs one-third to almost one-half the length of the corolla, rather thick, tapering, incurved; stamens about .5 cm. long; filaments linear, anthers ovate; capsule ovate, attenuate, about 1.5 cm. long. Distribution: p&ramos of Ecuador. Specimens examined: Ecuador: Sangai, Karslen (V); Azuay, Spruce 5181 (V); Quitensian Andes, coll. of 1855, Covlhouy (F); Farm of Antesiana, Nov. 2, 1858, Jameson (ANSP); same locality, alt. 5000 m., Oct. 1923, Anthony & Tate 299 (US); Rucu-Pichincha, Aug. 1923, Anthony & Tate 182 (US); Prov. Carchi, paramos 12 miles west of Tulcan, alt. 3300 m., Aug. 10, 1923, Hitchcock 20909 (G, NY, US); Chimborazo, Hans Meyer 118 (BG tyi-e). 58. H. barbicaulis Gilg in Engl. Bot. Jahrb. 54: Beibl. 118, p. 112. 1916. Annual herb, up to 2 dm. high; flowering stem seemingly erect, simple below, frequently branched above, branches erect, about .2 cm. thick, densely leafy, 4-alate, densely, long-ciliate below each node; internodes 2-4 cm. long; cauline leaves herbaceous, narrowed into broad ciliolate petioles, oblong, up to 2 cm. long, .5-6 cm. broad, acute or very acute, subapiculate, 5-nerved, veins inconspicuous above, prominent beneath, loosely reticulate; inflorescence a terminal dense many-flowered (7-9) subumbellate cyme, or 3-f lowered axillary cymes; pedicels of apical flowers 3-4 cm. long, lateral 2 cm. or less, all winged; calyx-lobes obovate- oblong, .6-7 cm. long, .35-4 cm. broad, apiculate, obsoletely 3- nerved; corolla about 1 cm. long, tube over one-third the length of the entire corolla; lobes ovate, subrotund, erose; spurs almost 1933J ALLEN — THE GENUS HALENIA 199 one-half the length of the corolla, pendulous, divaricate; stamens .2-3 cm. long; anthers ovate-oblong. Distribution: Peru. Specimens examined: Peru: Chacapoyas, Dec. 1846, Mathews (DH type). 59. H. Rusbyi Gilg in Engl. Bot. Jahrb. 54: Beibl. 118, p. 117. 1916. Perennial herb up to 3 dm. high; root strong, thick, ligneous, frequently horizontal, curved at the apex, sending up one or more simple erect flowering stems, bearing marcescent leaves at the base, or short densely leafy sterile branches ; basal leaves slightly narrowed into broad petioles as long as the leaf blades, elliptic, frequently up to 3 cm. long, acute, 3-nerved; cauline leaves 1-4 pairs, becoming sessile toward the summit of the stem, up to 3 cm. long, oblong-lanceolate, attenuate, acuminate; inflorescence terminal and axillary many-flowered umbellif orm cymes ; pedicels erect, up to 2 cm. long; calyx-lobes narrowly ovate, up to .7 cm. long, acute, 3-nerved; corolla up to 1.4 cm. long, tube .25-.3 cm. long; lobes ovate, acute; spurs approximately one-half or slightly more than one-half the length of the corolla, almost horizontally divergent, often with the tips incurved; stamens about .35 cm. long; filaments linear, anthers broadly ovate; capsule up to 1.5 cm. long, ovate-lanceolate, subfalcate; seeds ovoid. Distribution: Bolivia. Specimens examined: Bolivia: Unduavi, alt. 3300 m., Oct. 1885, Rusby 670 (ANSP, G, M F NY, US, V type). Similar to H. Purdieana, but a sturdier plant with broader flowers, spurs more divergent and more strongly recurved at tip, stem thicker, and leaves larger. 60. H. Purdieana Wedd. Chlor. And. 2: 76. pi. 58 A. 1859. Perennial herb, up to 2.5 dm. high; root ligneous; sterile stems 1-3, short, leafy; fertile stems probably more than one, simple, erect, slender; basal leaves in dense rosette, attenuate into slender petioles nearly equalling the blades in length, lanceolate to linear, up to 1.5 cm. long, .3 cm. broad, acute, obscurely 3-nerved, mid- vein prominent; cauline leaves 3-5 pairs, at intervals of about 4 cm., more or less closely appressed, practically sessile, linear, [Vol. 20 200 ANNALS OF THE MISSOURI BOTANICAL GARDEN attenuate, 2 cm. or less long, acuminate; inflorescence a terminal, rarely axillary as well, 1-5-f lowered cyme; pedicels up to 2.5 cm, long, recurving conspicuously at tip; calyx-lobes lanceolate, up to .7 cm. long, attenuately acuminate, 3-nerved; corolla up to 1.8 cm. long, greenish-white (fide Killip & Smith 18665, corolla-tube and lobes greenish-white, spurs while; Linden 729, fl: blanches), tube less than one-third the length of the entire corolla; lobes ovate, acute, erose; spurs nearly one-half the length of the corolla, slender, tapering, divergent, but slightly depressed at tip; capsule approximately 1.4 cm. long, ovate, attenuate at tip. Distribution: Colombia. Specimens examined: Colombia: Prov. de Pamplona, Pdramo de las Cruces, alt. 3000 m., Nov. 1842, Linden 729 pp. (BG TTP1B, DII, M photo, V); "Dept. Norte de Santander: between Mutiscua and Pamplona," alt. 3400 m., Feb. 23, 1927, Killip & Smith 19723 (US); "Dept. Norte de Santander: Pilramo de Romeral," alt. 3800-4200 m., Jan. 30, 1927, Killip & Smith 18665 (US); "Dept. Santander: Pdramo de las Puentes," above La Baja, alt. 3500-3700 m., Jan. 25, 1927, Killip d- Smith 18229 (US); "Dept. Santander: Pdramo de Romeral," alt. 3800-4000 m., Jan. 29-30, 1927, Killip & Smith 18546 (NY, US); same locality, 3800-4200 m., Jan. 80, 1927, Killip & Smith 186U (US); same locality and date, Killip & Smith 18688 (US); Pdramo de las Coloradas, above La Baja, alt. 390O-4100 m., Jan. 27, 1927, Killip & Smith 18426 (M, NY, US); same locality and date, Killip & Smith 18466 (NY, US); western slope of Pdramo Rico, alt. 36(K) m., Jan. 15-19, 1927, Killip & Smith 17722 (US). The majority of the specimens collected by Killip & Smith vary from the typical members of the species only in that the spurs are slightly more pendulous. The habit is similar. This differ- ence is not considered of varietal importance, hence the specimens are placed in the species proper. The species H. Purdieana has a marked resemblance in habit and in the color of the flower to H . Pringlei of Mexico, but the spurs are not reflexed to the extent they are in the former. 60a. H. Purdieana var. congesta* 5 Allen, n. var. Plant shorter than the species, not more than 1 dm., usually less than .5 dm., high; root heavy, extremely ligneous; stems 1- 66 H. Purdieana var. congesta Allen, var. nov. — Planta speciei breviora, rion plusquam 1 dm. alta, plerumque minusquam .5 dm. alta; radice crassa, lignissima; caul i bus 1-multis, erectis, simplicibus; foliis basalibus speciei pluribus, in rosula densissima, subcoriaceis, linearibus vel lineari-lanceolatis, minusquam 1.5 cm. longis; foliis caulinis 1-2 geminis, lineari-lanceolatis, adpressis; inflorescentia speciei simili. Colomiua: Dept. Santander, PAramo de Santurb&n, near Vetas, alt. 3950-4160 m., Jan. 17, 1927, Killip & Smith 17568 (M type, NY, US). 1933] ALLEN — THE GENUS HALENIA 201 many, erect, simple; basal leaves more numerous than in the species, and in very dense rosettes, subcoriaceous, linear to linear-lanceolate, less than 1.5 cm. long; cauline leaves 1-2 pairs, linear-lanceolate, very closely appressed; inflorescence similar to that of species. Distribution: known only from the Dept. of Santander, Colombia. Specimens examined: Colombia: "Dept. Santander: Paramo de Santurban," near Vetas, alt. 3950-4160 m., Jan. 17, 1927, Killip & Smith 17568 (M type, NY, US); same locality and date, Killip & Smith 17485 (NY, US); Paramo Frailejonale, near Vetas, alt. 3750-3850 m., Jan. 21, 1927, Killip & Smith 17982 (NY, US); Paramo de Mogotocoro, near Vetas, alt. 3700-3800 m., Jan. 18, 1927, Killip & Smith 17604 (NY, US); Paramo Rico, near Vetas, alt. 3750-3850 m., Jan. 18, 1927, Killip & Smith 17662 (M, NY, US); Paramo de las Vegas, alt. 3700-3800 m., Dec. 20-21, 1926, Killip & Smith 15680 (US) ; Paramo de Santurban, en route from Tona to Mutiscua, alt. 3800-4300 m., Feb. 18, 1927, Killip & Smith 19551 (US). 61. H. Hieronymi Gilg in Fedde, Rep. Spec. Nov. 2: 52. 1906. Annual, up to 3.5 dm. high; stem single, simple, erect; basal leaves few, in a rather loose rosette, petiolate, oblanceolate, up to 2.5 cm. long, .25 cm. broad; cauline leaves 4-5 pairs, membran- aceous, sessile, lanceolate, 1.5-5 cm. long, .3-.9 cm. broad, acute, obsoletely 3-nerved; inflorescence a terminal 5-6-flowered sub- umbelliform cyme, or often solitary axillary flowers, pedicels up to .7 cm. long; calyx-lobes oblanceolate, .6-.7 cm. long, .2 cm. broad, acute; corolla about 1 cm. long, yellow-green; lobes ovate, acute; spurs nearly one-half the length of the corolla, pendulous, slightly spreading. Distribution: Argentina. No specimens examined, but description compiled from original publication and photograph, (type, Fiebrig 261$, BG). 62. H. bifida 66 Rusby & Allen, n. sp. Perennial, up to 3 dm. high; root thick, ligneous; stem single, simple, erect, rather stiff, stout, winged; basal leaves in a dense 66 H. bifida Rusby & Allen, sp. no v. — Herba perennis, usque ad 3 dm. alta; radice crassa, lignea; caule solitario, simplice, erecto, paulo rigido, robusto, alato; foliis basalibus, in rosula densa, rigidis, petiolatis, oblanceolatis, usque ad 4.5 cm. longis, .4 cm. latis, acutis vel subito acuminatis, 3-nerviis; foliis caulinis 3-4 geminis, ses- silibus, lanceolatis, 2-3 cm. longis, decrescentibus sursum, .3 cm. latis, acuminatis, 3-nerviis, medio-nervo infra prominenti, supra immerso; inflorescentia multo-flor- [Vol. 20 202 ANNALS OF THE MISSOURI BOTANICAL GARDEN rosette, stiff, petiolate, oblanceolate, up to 4.5 cm. long, .4 cm. broad, acute to abruptly acuminate, 3-nerved; cauline leaves 3-4 pairs, sessile, lanceolate, 2-3 cm. long, decreasing toward the summit, .3 cm. broad, acuminate, 3-nerved, in id vein prominent below, sunken above; inflorescence a many-flowered terminal subumbellate cyme, frequently fewer-flowered cymes in axils of upper leaves; pedicels up to 3 cm. long; calyx-lobes broadly oblanceolate, .5 cm. long, .15 cm. broad, acute, obsoletely 3- nerved; corolla up to 1.3 cm. long, tube one-third the length of the entire corolla; lobes ovate, more or less acute, apiculate, erose; spurs more than y 2 the length of the entire corolla, slender, tapering, pendulous, spreading, incurved at tip; stamens .25 cm. long; filaments linear; seeds ovoid, dark brown. Distribution: Bolivia. Specimens examined: Bolivia: Cocopunco, alt. about 3000 m., March 24-30, 1926, Tate 879 (NY type). 63. H. Weddelliana Gilg in Engl. Bot. Jahrb. 25: 724. 1898; 54:Beibl. 118, p. 118. 1916. H. antigonorrhoica Gilg in Engl. Bot. Jahrb. 54: Beibl. 118, p. 119. 1016. Perennial herb from 1.5-3 dm. high; root fibrous; subterranean stem more or less elongate, densely covered with remains of marcescent leaves, horizontal, more or less erect, many-stemmed; sterile stems leafy, numerous; flowering stems numerous, curved or erect, the lower part densely leafy, the upper loosely leafy, internodes 2-8 cm. long; basal leaves numerous, in rosette, narrowly petiolate, subconnate, spatulate to oblong-lanceolate, 1-3 (rarely 5) cm. long, .2-.6 cm. broad, acute, 3-nerved; cauline leaves subconnate, lanceolate to oblong-lanceolate, 1-2 (rarely 6) cm. long, .2-6 cm. broad, acute, 3-nerved; inflorescence usually terminal, occasionally lateral, subumbellate or subcapitate many- ifera, terminali, subumbellato-cymosa; saepe cymis pauciore-floriferis in superiorum foliorum axibus; pedicellis usque ad 3 era. longis; calycis lobis late oblanceolatis, acutis, obsolete 3-nerviis, ca. .5 cm. longis, .15 cm. latis; corolla usque ad 1.3 cm. longa; tubo \^ totae corollae longitudini adacquanti; lobis ovatis, plusminusve acutis, apiculatis, erosis; calcaribus tenuibus, plusquam K totae corollae longitu- dini adaequanti, attenuatis, pendulis, divaricatis, ad apicem incurvatis; staminibus .25 cm. longis; filamentis linearibus; seminibus ovoideis, brunneis. — Bolivia: Coco- punco, alt. about 3000 m., March 24-30, 1926, Tate 879 (NY type). 1933] ALLEN — THE GENUS HALENIA 203 flowered cymes; pedicels .3-8 cm. long, the central one longer; calyx-lobes obovate-oblong to oblong, .5-8 cm. long, acuminate or acute, obsoletely 3-nerved; corolla 1.2-1.5 cm. long, yellow- green, tube less than one-half the length of the entire corolla; lobes ovate, acute to subrotund, entire or crenulate, erose; spurs one-half (rarely three-fourths) the length of the corolla, subhori- zontally divaricate but incurved at the apex; stamens .25-.35 cm. long; filaments linear usually, anthers ovate; capsule lanceo- late-ovate. Distribution: mountains of Colombia, Ecuador, and Peru. Specimens examined: Colombia: near Pasto, alt. 3600 m., June 13, 1878, Lehmann (V); p&ramos of Guanacas, Prov. Popayan, Lehmann 809 (NY); same locality, Lehmann 6128 (BG type, F, US); same locality, Hartweg 1255 (NY, V type of H. antigonorrhoica Gilg); grassy p&ramo, Paletara, Dept. of Cauca, June 15-17, 1922, Pennell 6929 (ANSP, Quito, alt. 4000 Lehmann 3498 Andes, alt. 4000 m., March 1859, Jameson (NY, US, V); 1859, Jameson (BG G, M photo, UC); without locality or date, Jameson (NY, US); Quitian Andes, Jameson (V); Andes, Quito, 1855, Couthouy (ANSP, G, NY); same locality, 1857-9, Spruce 5131 (G, UC, V); San Ignacio, Pichincha Region, alt. approximately 4000 m., Aug. 14-19, 1923, Anthony & Tate 133 (US); Pichincha, alt. 3500 m., Feb. 3, 1927, Firmin 5 (US); Pichincha-Quito, Karsten (V); same locality, Dec. 30, 1929, HeUborn 137 (S); Tunguragua, above Bafios, alt. 2000 m., Feb. 27, 1920, Holmgren 378 (S); Cotopaxi, alt. 3000-4000 m., Dec. 27, 1879, Lehmann (V); without locality, Gesner t~* • i 7 -4 F* t / y Tn m r\ ft I « • i • « i« 4200 2800-4000 Peru: grassy places in shrub-wood, Putis, Choimacota Valley, Prov. Huanta, 3400-3500 10-24. 1923. Macbride 64. H. elegans 67 Allen, n. sp. Stout perennial, up to 4 dm. high; root thick, fibrous; axis covered with blackened remains of leaf bases, erect, sometimes 67 H. elegans Allen, sp. nov. — Herba perennis, usque ad 4 dm. alta; radice crassa, fibrata; axe foliorum reliquiis tecto, erecto, saepe plus minusve decumbenti; ramis brevibus sterilibus, dense foliosis, 10-12 cm. altis; caule solitario, florifero, crasso, .5-6 cm. in diametro, infra simp lice, supra parum ramoso, striato; foliis basalibus in rosula densa, petiolatis, late lanceolatis, 7-9 cm. longis, acuminatis, prominenti 3-nerviis, nervis supra immersis, infra prominenti; foliis caulinis 4-6 geminis, ses- silibus, 2-4 cm. longis, sursum decrescentibus; inflorescentia cymosa, laxa, multo- florifera, terminali vel ramis brevibus clause adpressis terminatis, laxis spicis similibus; pedicellis usque ad 4 cm. longis, ad apicem parum curvatis; calycis lobis foliosis, late lanceolato-ellipticis vel ovatis, 1-1.2 cm. longis, .4 cm. latis, attenuate acuminatis, prominenti 3-nerviis; corolla 3-4 cm. longa, viride, tubo y$- x /i corollae longitudini adaequanti; lobis obovato-ovatis, erosis, apiculatis; calcaribus prope [Vol. 20 204 ANNALS OF THE MISSOURI BOTANICAL GARDEN more or less decumbent; short sterile branches, densely leafy, 10-12 cm. high; single flowering stem, coarse, .5-6 cm. in di- ameter, simple below, slightly branched above, striate; basal leaves in dense rosettes, petiolate, broadly lanceolate, 7-9 cm. long, acuminate, prominently 3-nerved, nerves sunken above, prominent below; cauline leaves 4-6 pairs, sessile, 2-4 cm. long, decreasing toward the summit; inflorescence loose, many-flowered, terminal cymes, or cymes terminating the short, closely appressed branches, giving the inflorescence a loose, spike-like appearance; pedicels up to 4 cm. long, slightly curved at the tip; calyx-lobes foliaceous, broadly lanceolate-elliptic or ovate, 1-1.2 cm. long, .4 cm. broad, attenuate-acuminate, prominently 3-nerved; corolla 3-4 cm. long, green, tube one-sixth to one-fourth the length of the entire corolla; lobes obovate-ovate, erose, apiculate; spurs nearly one-half the length of the corolla, slender, with tips extremely glandular, pendulous, curved outward, somewhat spreading, approximate at base; stamens about 1 cm. long; filaments linear, anthers narrowly ovate; capsule attenuate-ovate. Distribution: Colombia. Specimens examined: Colombia: Dept. Santander, Paramo de Romeral, alt. 3800-4100 m., Jan. 29-30, 1927, Killip & Smith 18668 (M type, NY, US). 65. H. Hoppii Reimers in Engl. Bot. Jahrb. 62 : 335. 1929. Perennial about 1 dm. high; flowering stem simple, erect, rather stout ; basal leaves in a dense rosette, herbaceous, petiolate, oblance- olate, 3 cm. long, .4 cm. broad, acute, nerved; cauline leaves 1-2 pairs, similar to a rosette, but petioles shorter; inflorescence atermi- nal, many-flowered cyme; pedicels 1-3 cm. or more long; calyx-lobes ligulate to elliptic, .6 cm. long, .2 cm. broad, acute at apex, sub- apiculate, obsoletely 3-nerved; corolla 1 cm. or less long, yellow- green, tube about one-third the length of the entire corolla; lobes acutish ovate, subrotund; spurs very slender, one-third to one- half the length of the corolla, pendulous, divergent at apex; stamens .35 cm. long. corollae longitudini aelaequanti, tenuibus, ad Hpicem glandulosis, pendulis, curvatia cxtrinseeis divergentioribus ad basin approximatis; staminibus ca. 1 cm. longis; filamentiglinearibus; antheris attenuate ovatis; capsula attnmata-ovata. — Colombia: Dept. Suiitander, paramo de Romeral, alt. 3800-4100 in., Jan. 29-30, 1927, Killip & Smith 18668 (M type, NY, US). 1933] ALLEN — THE GENUS HALENIA 205 Distribution: Colombia and Peru. Specimens examined: Colombia: (type not seen, Hopp 33a BG, M photo). Peru: Cuzco, alt. 3000-3600 m., July 1923, Herrera (US). 66. H. asclepiadea (HBK.) G. Don, Gen. Hist. 4: 177. 1838; Griseb. Gen. & Sp. Gent. 326. 1839; in DC. Prodr. 9: 129. 1845 pp ; in Linnaea 22 : 45. 1849 ; Benth. PI. Hartw. 228. 1846 ; Wedd. Chlor. And. 2: 75. 1859, pp. Swertia asclepiadea HBK. Nov. Gen. & Sp. PI. 3: 175. 1818. S. quadricornis Willd. ex Roem. & Schult. Syst. Veg. 6: 134. 1820; fide Gilg, I c. Perennial herb, up to 4 dm. high; fertile stems one or more, simple, erect; one to several short densely leafy sterile stems arising from the root or, rarely, from the base of a flowering stem ; basal leaves attenuate into a more or less slender elongate petiole, narrowly lanceolate, 3-4 cm. long, up to .5 cm. broad, attenuate- acuminate, 3-nerved; cauline leaves sessile, more elongate than the basal leaves but otherwise similar to them; inflorescence a terminal or axillary, 3-8-flowered, occasionally umbelliform cyme, petioles up to 3.5 cm. long; calyx-lobes broadly lanceolate, up to .6 cm. long, acute, 3-nerved; corolla approximately 1.2-1.3 cm. long, greenish-yellow, tube one-fourth the length of the entire corolla; lobes broadly ovate, acute; spurs horizontal, measuring up to 1.7 cm. from tip to tip; stamens approximately .45 cm. long; filaments linear, anthers ovate; capsule up to 1.8 cm. long, ovate, attenuate at tip, subfalcate; seeds oval-elliptic, reticulate. Distribution: Colombia, Ecuador, and Peru. Specimens examined: Colombia: Dept. of Cundinamarca, above Bogota, bushy slope, alt. 2700-2800 m., Aug. 16, 1917, Rusby & Pennell 1269 (NY); Paramo de Guasca, July 21, 1919, Bro. Ariste- J oseph AS90 (US); mountains near Bogota, Oct. 20, 1852, Holton 468 (ANSP, DH, G, NY); paramos, Bogota, March 1916, M. T. Daire 123 (US); Bogota, alt. 2900 m., Karsten (V); Bogota, alt. 3000 m., 1851-57, Triana 1958 (V); near Bogota, Goudot (DH); Andes near Bogota, 1843, Hartweg 1254 (DH, NY, V); same locality, Bro. Ariste- J oseph (US); Pamplona, Paramo de las Cruces, alt. 3600 m., Nov. 1842, Linden 729 pp. (DH, V); Tequendama, 1917, Bro. Ariste- J oseph A46 (G, US); paramos, Guasca, alt. 4000 m., Aug. 1931, Arbelaez 1201 (US); (type not seen, Humboldt & Bonpland, HJP, M, photo). Ecuador: Prov. Loja, between San Lucas and Ofia, alt. 2200-3100 m., Sept. 7, 1923, Hitchcock 21539 (G, NY, US). Peru: Dept. of Cusco, Paso de Tres Cruces, Cerro de Cusilluyoc, alt. 3800-3900 m., May 3, 1925, Pennell 13842 pp. (NY). 206 ANNALS OF THE MISSOUKI BOTANICAL GARDEN [Vol. 20 This species is closely related to H. recurva of southwestern United States and Mexico. 67. H. Kalbreyeri Gilg in Engl. Bot. Jahrb. 54: Beibl. 118, p. 120. 1916. Perennial caespitose herb up to 2.5—4 dm. high; axis procum- bent, slender, subligneous, the upper part erect; stems 1-several, erect, more or less slender, simple, internodes 3.5 cm. long; sterile stems up to 7-8 cm. high, densely leafy above; rosette leaves petiolate, herbaceous, stiffly erect, never drooping, ob- lanceolate, about 5 cm. long, .3 cm. broad, acute, 3-nerved, midvein heavy and hard ; cauline leaves often as many as 8 pairs, sessile, linear-lanceolate to lanceolate, up to 2.5 cm. long, .2-.25 cm. broad, acute, 3-nerved; inflorescence terminal or axillary cymes, 1-5-flowered, frequently pseudo-spicate or pseudo- paniculate, pedicels up to 3 cm. long; calyx-lobes lanceolate, up to .5 cm. long, .15 cm. broad, 3-nerved; corolla 1.2-1.5 tube one-third the length of the entire corolla; lobes ovate, some- what acute, erose; spurs one-half to three-fourths the length of the corolla; stamens about .4 cm. long; filaments linear, anthers broadly ovate; capsule lanceolate. Distribution: Colombia. Specimens examined: cm 3700 & Bro. AHsle-J oseph 11924 (ANSP, US); Dept. Cusco, La Raya, April 22, 1925, Pennell 18518 (ANSP); Dept. Norte de Santander, between Mutiscua and Pamplona, alt. 3400 m., Feb. 23, 1927, Killip & Smith 19728 (US), (type not seen, Kalbreyer 717, BG, M photo). 68. H. bella Gilg in Fedde, Rep. Spec. Nov. 2: 52. 1906. Perennial herb, 2-2.5 dm. high; stems decumbent, bearing al apex numerous short sterile leafy branches up to 10 cm. long and elongate fertile branches, simple, erect, more or less slender internodes up to cm. long; basal leaves numerous, subcoriaceous petiolate, oblanceolate to lanceolate, 3-3.5 cm. long, .4 cm broad, acute, 3-5-nerved, veins sunken above, prominent below of sterile branches 3-4 pairs, sessile, lanceolate, acute; inflorescence a terminal sub- umbelliform many-flowered cyme, rarely axillary 3-flowered cymes; calyx-lobes obovate-lanceolate, .6-.7 cm. long, .2 cm. 1933] ALLEN — THE GENUS HALENIA 207 broad, acute, nerved; corolla approximately 1.6 cm. long, green; lobes obovate, acute; spurs horizontal or subhorizontal, about three-fourths the length of the corolla. Distribution: Peru. Specimens examined: Peru: Dept. Junfn, Prov. Tarma, Huacapistana, alt. 3000-3100 m., Weberbauer 2065 (BG type, DH, M photo); Pavon 737 (DH). 69. H. sphagnicola Gilg in Engl. Bot. Jahrb. 54: Beibl. 118, p. 121. 1916. Perennial, about 2.5 dm. high; root slender, multifibrous ; axis short, slender, nearly erect; several curved-ascending sterile branches; fertile branches single, simple or branched at base, erect, the lower portion densely leafy, the upper loosely leafy; basal leaves in rosette, herbaceous, attenuate into petioles once or twice the length of the blades, obovate, 1.5-2.2 cm. long, .5 cm. broad; lower cauline leaves several pairs, petiolate, obovate, 1.5-2 cm. long, .3 cm. broad, acute; upper cauline leaves slightly narrowed at base but more or less sessile, oblong or oblong- lanceolate, .7-1.2 cm. long, .2-25 cm. broad, acute to acuminate, obsoletely 3-nerved; inflorescence a terminal many-flowered sub- globose cyme; pedicels about 1-1.3 cm. long; calyx-lobes obovate to ovobate-oblong, scarcely .3 cm. long, .1-.15 cm. broad, sub- rotund at apex, obsoletely 3-nerved; corolla .8-9 cm. long, sulphur-yellow, tube less than one-half the length of the entire corolla; lobes ovate, rotund; spurs slender, tapering, one-half the length of the corolla, but subhorizon tally divaricate, often curved upward at the apex. Distribution: Peru. No specimens examined, but description compiled from original publication and photograph, (type, Weberbauer 4376, BG, M, photo). List of Exsiccatae The distribution numbers are printed in italics. Unnumbered collections are indicated by a dash. The numbers in parentheses are the species numbers used in this monograph. Aiton, G. B. — (23). Andrieux, G. 226 (31). Alaman, L. — (2a). Anthony, H. E. & Tate, G. H. H. 133 Allen, J. A. — (23). (63); 182, 299 (57). Altamirano, F. 908 (21). Arbelaez, E. P. 1201 (66). 208 [Vol. 20 ANNALS OF THE MISSOURI BOTANICAL GARDEN Archer, W. A. 1348 (2). Aristc-JoHcph, Hro. — , A46 (66); A42S (10); AS90 (66). Arnold, I. S. Arsdne, Hro. G. (23). (2a); (2d); Eggleston, W. W. — , 3051 (23); 1077 A, 15781, 17137 (22). Eggleston, W. W. & Fernald, M. L. (23). Ehrenberg, K. 608 (30). (31a); 37 (23); 55 (2d); 1703, 1711 Ellin, C. C. 20 (22). (2a); 2098 (2); 5610 (2a); 5864, 5957 Endlich, R. 53 (22). (2d). Bailey, W. W. — Bang, M. 666 (45); 1019 (48). Farr, E. M. (23). (23). Farwell, O. A. 249, 770 (23). Banks, J. (23); (23a). Faxon, C. E. Fellows, D. W. (23). (23). Bergman, H. F. 2948 (23). Berlandier, J. L. 1207 (2, 2a). Beyrich, II. K. (2a). Blake, J. (23). Blumer, J. C. 1359 (22). Boott, W. (23). Bourgeau, E. — (23); 799, 3126 (2a). Bowdoin College, 290 (23a). Boyce, T. E. (23). Brenton, M. 10.— (23); 144a f (23a). Brinkman, A. H. 3678 (23). Brown, S. 667 (23). Buchtien, O. 54, 600 (55); 1484 (35). Burglehaus, P.— (23). Canby, W. M. Chapman, A. W. Churchill, J. R. Chute, A. P. — (23). (23). (23). - (23). Collins, J. F., Fernald, M. L. & Pease, A.S. — , 4261 (23). Conzatti, C. 2295, 4265 (26). Conzatti, C. & Gonzalez, V. 878 (2b). Cook, O. F. 45 (32a). Cook, O. F. & Gilbert, G. B. 1808 (56); Fried richstahl, E. Fernald, M. L. — , 87, 1151 (23). Fernald, M. L. & Collins, J. F. 244 (23). Fernald, M. L., Dodge, C. W. & Smith, L. B. 25986 (23). Fernald, M. L. & Long, B. 14390, 14392 (23); 28950 (23a). Fernald, M. L., Long, B. & Dunbar, B. H. 26982, 26983, 26984, 26985, 26986 (23a). Fernald, M. L., Long, B. & Fogg, J. M. 881 , 382 (23a) . Fernald, M. L. & Pease, A. S. 16624 (23). Fernald, M. L. & Wiegand, K. M. 3908, 8909, 6081 (23a) . Fernald, M. L., Wiegand, K. M. & Kittredge, J. 8910 (23a); 3911 (23); 3912 (23a). Fernald, M. L., Wiegand, K. M. & Long , B. 28951 (23a). Firmin, Fr. G. 5 (63); 697 (42). Forbes, F. V. Fowler, J. — (23). - (23). Franklin & Douglas (23). (24). 1793 (36); 1811 (56). Coulter, T. 939 (31). Couthouy, J. P. (5i); (57); (63). Cristan, F. (24). Cumming, H. 128 (55). Daire, M. T. 123 (66). Dodge, C. K. 852 (23). Dombey, J. Don, D. — (56). (31). (23). Drexter, C. Drummond, J. Eames, E. A. Eames, E. H. & Godfrey, C. C. 8030 Goderiels, Funck, N. 415 (15). Funck, N. & Schlim, L. J. 901 (7); 11 48 (8). Galeotti, H. 1488 (30); 1489 (26); 1490 (2b); 7166 (33); 7219 (2); 7220 (30); 7221 (1); 7222 (31). Garcia, P. Ibana — 410 (25). Gates, F. C. 14156 (23). Gates, F. C. & M. T. 9768, 10716 (23). Gehiiger, W. 92a (8). - (23). (23). Gesner, (63). Ghiesbreght, A. 137, 618 (2). (23). (23a). Goodale, G. L. (23). 1933] ALLEN — THE GENUS HALENIA 209 Goudot, (10); (66). Karsten, H. (5); (10); (42); Greenman, J. M. & M. T. 5994 (24b). (57); (63); (66). Grisebach, A. H. R. — (2) ; Gutzwiller, 82 (18). Haberer, J. V. — , 601 (23). Hadley, J. 1 (23). (23). Karwinsky, W. F. von, 122 (31). Kennedy, G. G. (23). Hall, C. H. (23). Hallowell, S. M. Harper, E. T. — (23). - (23). Harshberger, J. W. 137 (2a, 30). Hartweg, T. 210 (2b); 847 (31a); 494 (33) ; 1254 (66) ; 1255 (63). Hasse, H. E. Hay, G. U. Haydon, W. - (23) . (23). (23). Heilborn, O. 137 (63); 437 (42). Heinburger, (23). Heller, A. A. 391 (31a); 401 (2). Kennedy, Mrs. J. J. 38 (23). Killip, E. P. 6889 (2). Killip, E. P. & Ariste-Joseph, Bro. 11924 (67). Killip, E. P. & Smith, A. C. 15679 (3) ; 15680, 17485 (60a); 17516, 17621, 1 7666 (50) ; 1 7568, 1 7604, 1 7662 (60a) ; 17722 (60); 17982 (60a); 18229, 18426, 18466, 18546 (60); 18568 (64); 18644, 18665, 18688 (60); 19551 (60a); 19723 (60, 67); 21737 (44); 22087 (43). Kittredge, E. M. Kofoid, C. A. (23). Herrera, F. L. (65). (23). Kuntze, O. 2356 (24); 23788 (2). Herzog, T. 2377 (54) ; 2402 (55) . Heyde, E. T. & Lux, E. 4729 (34) . Hill, E. J. 179 (23). Lange, J. (23). Lankester, C. H. 670 (24). Lechler, W. 2001 (56). Hitchcock, A. S. 20909 (57); 21539 (66). Lehmann, F. C. — (66); 809, 3498 (63); Hoffmann, C. 119 (24b). Holmgren, I. 378 (63). 4878 (49); 6128 (63); 6190 (12); 7860 (17). Holton, I. F. 19 (464) (2); 467 (10); 468 Lemmon, J. G. (2a); (66). Holway, E. D. W. Lemmon, J. G. & wife (22). (2a); (22). (23). Leston, (23). (23). (23). Hooker, J. D. - Hosford, F. H. Houghton, D. 34 (23). House, H. D. 5643, 16688 (23). Howe, C. D. (23a) . Liebmann, F. M. 10770 (33); 10771 (26); 10774 (30); 10775 (2); 10776 (30); 10777 (2c); 10778 (2); 10780 (31). Linden, J. 437 (6) ; 456 (2) ; 729 (60) ; 729 (66); 934 (\))985 (31a). Howe, C. D. & Lang, W. F. 1298, 1403 Loew, O. (23a). Humboldt, A. & Bonpland, A. Loring, (22). (23a) ; (23). (2) ; Louis-Arsene, Fr. 365, 403 (23a) . (31); (42). Lour, O. (22). Hyatt, A., Shaler, N. & Verrill, A. E. Macbride, J. F. 3040 (47); 8072 (36); (23). Jahn, A. 558, 839 (7). Jameson, W. (49); Jones, M. E. (22). (2); 53 (40); (42); 3431 (2); 4362 (63). Macbride, J. F. & Featherstone, 898 (38); 1 657, 2001 (56). (57); (63). Macfarlane, (23). (2e); (25); Mackenzie, K. K. 3618 (23). Mackenzie, K. K. & Griscom, L. 10411, Julio, Bro. 838, 455 (55). 11135 (23a). Jurgensen, 386 (33); 812 (26); 811 MacNab, W. (23). (30). Kalenborn, A. S. 91 (38). Kalenborn, M. 91 (38). Macoun, J. — , 77, 300, 1191, 54300 f, 68784 f (23) . Mandon, G. 369 (36, 52); 369 (56). 210 [Vol 20 ANNALS OF THE MISSOURI BOTANICAL GARDEN jnnell, F. W. & Ha: 9894 (16); 9997 (14). Marie-Victorin, Bro. 28, 8849 (23). I Marie-Victorin, Bro., Brunei, J. B., Rolland-Gcrmaine & Rousseau, Z. Penncll, F. W 17668 (23). Pereival, Mrs. Marie-Victorin, Bro., & Rolland-Ger- Pickering, C. (23). (23). maine, 96 47, 27159 (23). 9881, 18482 (23a); Pinkerton, M. E. & Allen, C. K. (23). Marie-Vietorin, Bro., Rolland-Germaine & Louis-Marie, 21076 (23). Pittier, H. 39 (32); 744 (24); 1111 (14); 2975, 8499 (24); 13076 (24b). Pittier, II. & Tonduz, A. 10805 (24). Martin, (23). Mathew, G. V. Mathews, A. (2). (23). Poggenburg, J. F. (23). Porter, T. C. (58); 523 (44); 8133 Priest, M. E. Pringle, C. A. (23). (23a). (2e) ; (31a); (22); Maxon, W. R. & Hay, R. 3676 (28). Mell, P. I 1 . & Knopf, Metcalfe, O. B. 501 (22). Mexia, Y. 2693 (31). Millspaugh, C. F. 82 (23). Moore, A. H. 1211 (23). (23). Morison, (2:on8 178 deflexa 162 var. Brentoniana 167 Dombeyana 197 var. a 194, 196 var. brevieornis 182 elata 155 elegans 203 elong), Livingston and Tottingham (18), Van Alstine ('19), Meier and Halstead ('21), and by Tottingham and Rankin ('22); buckwheat, by Shive ('15, 17) and by Shive and Martin ('18, 18a); potato, by Johnston ('24) and by Martin and Shive ('20); celery, by Poole and Fant ('22); cranberry, by Addoms and Mounce ('31); peach seedlings, by Davidson ('28). After investigating the work of the above authors, it appeared to us that the triangular system of arranging a varying series of nutrient solutions provided the most satisfactory approach to the problem of the reaction of orchid seedlings to their inorganic substrate. Seeds for the investigation were furnished from the green- houses of the Missouri Botanical Garden through the courtesy of Dr. George T. Moore, the Director. Flowers of Cattleya Trianae Linden and Rchb. f. were pollinated November 17, 1930, and the largest and apparently the best pod was picked May 3, 1932, after a development period of almost eighteen months. It has 1933] WYND — ORCHID SEEDLINGS AND NUTRITIONAL IONS 225 been the author's experience that in general the capsules first to mature furnish seeds of lower vitality than those maturing later; hence the one requiring the longest time to mature was used to supply seeds for the present work. The nutrient solutions investigated were those of Type I and Type IV, as described by Livingston ('19). By the use of only three salts it is possible to arrange solutions containing the six major nutritional ions in six different combinations. Of these six possible combinations, only two will have all their ions added in unlike combinations. The compositions of the solutions used are given in tables i and n. In both types the total osmotic TABLE I TYPE I- -KH 2 PO\ Ca(NO')', MgSO* Culture Molecular proportion Volume-molecular concentration No. KH 2 PO< Ca(N0 3 ) 2 MgSO 4 KH 2 PO Ca(NO,) 2 MgSO* Rl SI 1 6 .0027 .0027 .0161 S2 2 5 .0025 .0049 .0123 S3 3 4 .0024 .0071 .0094 S4 4 3 .0022 .0089 .0067 S5 5 2 .0022 .0108 .0043 S6 6 1 .0020 .0122 .0020 R2 SI 2 1 5 .0053 .0027 .0132 S2 2 2 4 .0049 .0049 .0099 S3 2 3 3 .0047 .0071 .0071 S4 2 4 2 .0045 .0090 .0045 S5 2 5 1 .0041 .0104 .0021 R3 SI 3 1 4 .0076 .0025 .0101 S2 3 2 3 .0072 .0048 .0072 S3 3 3 2 .0068 .0068 .0045 S4 3 4 1 .0065 .0086 .0021 R4 SI 4 1 3 .0099 .0025 .0074 S2 4 2 2 .0094 .0047 .0047 S3 4 3 1 .0090 .0068 .0022 R5 SI 5 1 2 .0123 .0024 .0049 S2 5 2 1 .0118 .0047 .0023 R6 SI 6 1 1 .0145 .0024 .0024 226 [Vol. 20 ANNALS OF THE MISSOURI UOTANICAL C.ARDEN TABLE II TYPE IV— K,8C>4, CadliPOOt. Mg(NOi)i* Culture Molecvilar proportions Volume-molecular proportions no. K 2 S0 4 Ca(H 2 P0 4 )2* Mg(N0i)i K 2 S0< Ca(H 2 P0 4 ) 2 Mg(NO s ), Rl SI 82 S3 1 1 1 1 2 3 6 5 4 .0018 .0018 .0019 .0018 .0036 .0056 .0108 .0092 .0075 R2 SI 82 S3 2 2 2 1 2 3 5 4 3 .0037 . 0037 .0037 .0018 . 0037 . 0056 .0091 .0074 . 0056 R3 SI 82 83 3 3 3 1 2 3 4 3 2 .0056 .0056 . 0056 .0019 . 0037 .0056 . 0075 . 0056 .003S Rl SI S2 83 4 4 4 1 2 3 3 2 1 . 0076 .0077 . 0078 .0019 . 0039 . 0059 .0057 . 0039 .0019 R5 SI 82 5 5 1 2 2 1 . 0097 . 0098 .0019 . 0039 . 0039 .0019 R6 SI 6 1 1 .0116 .0019 .0019 * The .solutions containing the highest proportion of Ca(H2l'0«) 2 could not be pre- pared, because sterilization at the temperature and pll used caused precipitation. atmosph Mallinckrodt salts of high used. One-hundred-cc. portions of the placed flasks. 1.75 per cent of Merck's powdered "Reagent" agar added, and steri- lized by autoclaving at 20 lbs. pressure for 20 minutes. Follow- ing the work of La Garde ('29), who found that maltose was the best source of carbohydrate, a 2 per cent concentration of this sugar was used in all cultures. A study of the sugar der 10 per cent conditions hydrolyzed hydrolysis of this owed that about In addition to the six major nutrient ions there were added le-half part per million of mantranese as mantranese si i Inhale. and one-half part per million of boron as sodium borate ferric phosphate suspension was prepared as described by Ln A 1933) WYND — ORCHID SEEDLINGS AND NUTRITIONAL IONS 227 ston ('19) and 1 cc. added to each liter of nutrient solution. This gave ferric phosphate in the concentration of about 3 mgms. per liter. Owing to the sensitivity of orchids to the acidity of the medium, each solution was so adjusted that the pH after sterilization was 4.8 to 5.0. Tables in and iv indicate the pH relationships be- fore and after sterilization. The seeds were sterilized by shaking TABLE III THE ACIDITY RELATIONSHIPS OP TYPE I SOLUTIONS* A B c Sol. plus Fe, B, Mn, and 2% maltose pH as made up pH adjusted before autoclaving pH after 1.75% agar added and auto- claved Rl SI S2 S3 S4 S5 S6 5.74 5.10 4.93 4.90 4.83 5.08 3.93 3.91 3.90 3.90 3.90 3.90 4.9 5.0 5.0 5.0 5.0 5.1 R2 SI S2 S3 S4 S5 4.86 4.75 4.73 4.73 4.83 3.93 3.88 3.93 4.10 4.10 4.8 5.0 4.9 5.0 5.0 R3 SI S2 S3 S4 5.26 4.86 4.83 4.76 4.22 4.24 4.20 4.20 5.0 5.0 5.0 5.0 R4 SI S2 S3 4.83 4.81 4.75 4.20 4.22 4.24 5.0 5.0 4.9 R5 SI S2 4.86 4.83 4.24 4.22 4.9 4.9 R6 SI 4.80 4.29 4.9 * The pH determinations indicated in columns A and B were obtained by the quinhydrone electrode, while the results in Column C were obtained colorimetrically by the use of brom-cresol-green as an indicator. 228 [Vol. 20 ANNALS OF THE MISSOURI BOTANICAL GARDEN TABLE IV ACIDITY RELATIONSHIPS OF TYPE IV SOLUTIONS DETERMINATIONS MADE AS INDICATED IN TABLE III A B C Sol. plus Ke, H, Mn, and 2% maltose 1> 1 1 as made up pH adjusted before autoclaving pi! after 1.75% agar added and auto- claved Rl SI S2 S3 4.39 3.71 3.56 4.10 4.18 4.27 5.0 4.9 4.9 R2 SI S2 S3 4.10 3.67 3.59 3.89 4.20 4.29 5.0 4.9 4.9 R3 SI 82 S3 4.05 3.73 3.64 3.90 4.22 4.30 4.9 5.0 4.8 R4 SI S2 S3 3.91 3.69 3.54 3.91 4.25 4.32 5.0 4.9 4.9 R5 SI S2 4.34 3.78 3.81 4.15 4.9 5.0 R6 SI 4.27 3.90 5.0 minutes in a small vial of calcium hypochlorite pared as described by Wilson They were then inoculated directly nto the flasks of nutrient agar by a platinum needle. The flasks were incubated at 20-21 shaded greenhouse compartment. During the C summer days the greenh could be held below these effect. and but the short duration temperatures apparently was without harmful III. Results Type I. — The growth period of these cultures extended from April 15 until November 12, 1932. At the end of this time, the 1933] WYND — OKCHID SEEDLINGS AND NUTRITIONAL IONS 229 diameter of the seedlings was measured by a low-power microscope equipped with a calibrated eye-piece micrometer according to the method of Quednow ('30). The flasks had been very lightly inoculated so as to eliminate crowding of the growing seedlings. The figures in table v indicate the diameter in microns, each TABLE V GROWTH OF SEEDLINGS IN TYPE I SOLUTION. THE FIGURES REPRESENT DIAMETERS IN MICRONS Solution Series 1 Series 2 Rl SI 915 910 S2 1100 1130 S3 970 1020 S4 1015 1010 S5 925 940 S6 810 830 R2 SI (Contaminated) (Contaminated) S2 925 980 S3 855 845 S4 600 740 S5 740 690 R3 SI 805 775 S2 660 740 S3 690 725 S4 585 445 R4 SI 860 690 S2 620 620 S3 620 670 R5 SI 635 640 S2 475 585 R6 SI 680 615 based on an average of 25 individuals. Figure 1 shows graphi- cally the comparative development on the different media used. Figures 3 and 4 indicate the areas of the triangles corresponding to greatest, medium, and least growth. The circles are drawn to scale and represent comparatively the magnitude of the seedlings of the respective cultures. [Vol. 20 230 ANNALS OF THE MISSOURI BOTANICAL GARDEN Type IV. — The growth period of these cultures lasted from April 28 until November 20, 1932. Table vi presents the data for the growth on these solutions. As before, the diameter is indicated in microns and is the average of 25 determinations. Figure 2 shows graphically the comparative development. ?i3i i5c liS. 1* 3* Ji 34 llSS *4 3l Ht 34 ?|34 Pi 3* Hi 35 I 5f \ Hi 3% ton Hi 14 Rill *t3f IK 3* If 3l ii3^ a b Fig. 1. a: diagrammatic representation of the relative diameters of seedlings germinated on Type I solution. Series 1; b: relative diameters of seedlings germinated on Type I solution. Series 2. -**3t ftl5t liSi Pi3» 432 If St 1631 Pl33 1131 till Tfc3» 1431 *l3t 1431 *3*J *Z3J 14SJ 1l3S a b Fig. 2. a; diagrammatic representation of the relative diameters of seedlings germinated on Type IV solution. Series 1; b: relative diameters of seedlings germi- nated on Type IV solution. Series 2. Figures 5 and G show the areas of the triangles producing greatest, medium, and least growth, the circles being drawn as before to indicate the average diameters of the seedlings of the respective cultures. 1933] WYND — ORCHID SEEDLINGS AND NUTRITIONAL IONS 231 TABLE VI GROWTH OF SEEDLINGS IN TYPE IV SOLUTION. THE FIGURES REPRESENT DIAMETERS IN MICRONS Solution Series 1 Series 2 Rl SI 920 990 S2 825 825 S3 550 595 R2 SI 955 950 S2 740 740 S3 635 665 R3 SI 965 965 S2 765 770 S3 750 690 R4 SI 1010 950 S2 730 730 S3 570 630 R5 SI 975 1035 S2 715 715 R6 SI 675 705 IV. Discussion Examination of figs. 3 and 4 shows perfect agreement between the duplicate series of Type I in the areas of greatest growth. These areas are characterized by having very low concentrations of KH 2 P0 4 . The five best cultures in row 1 have only one-eighth of the total osmotic concentration due to this salt. The ratios of calcium nitrate to magnesium sulphate are seen to vary be- tween wide limits, .1667 to 2.500, with no significant effect on growth. The areas of medium and least growth do not show such perfect agreement in the duplicate series, but a comparison shows that the areas of medium growth correspond in general to those having the medium KH 2 P0 4 concentration, and the areas of least growth correspond to those having the highest proportion of this salt. This relation is particularly clearly shown in fig. 4. From these data one may be led to the conclusion that germinating orchid seeds and young seedlings are comparatively sensitive to 232 [Vol. 20 ANNALS OF THE MISSOURI BOTANICAL GARDEN varying concentrations of KH2PO4, but relatively insensitive to wide variations in Ca(N0 3 ) 2 and MgS0 4 . Since the ions were present as salts, it is impossible to determine which ion is responsible for the result. It is possible to add the six ions in but one other combination of three salts so that each anion will be linked with a different cation. This combination is represented by the Type-IV triangle. It is unfortunately impossible to prepare the complete Type-IV series because at the pH used (5.0) the solutions having the higher Ca(H 2 P0 4 )2 concentrations precipitated on sterilization due to the decom- position of this compound. Fig. 3. Type I, aeries 1, showing KH».P0t Fig. 4. Type I, series 2, showing areas of high, medium, and low yield. areas of high, medium, and low yield. Culture R2 SI omitted because of Culture R2 SI omitted because of As{>ergilliis contamination. Aspergillus contamination. Growth on the possible solutions of this type shows a remark- able result. The areas of greatest growth perfectly agree and are characterized by solutions having the minimal proportion of the total osmotic, concentration due to Ca(H 2 P04) 2 . Again, the ratios of the other salts, K 2 S0 4 to Mg (NC^ vary between wide limits, .1()(>7 to 2.500, with very little effect on growth. The areas of medium growth correspond to those of medium concen- tration of this salt, and those of least growth correspond to the areas of high concentration of this salt. This comparison of the growth on solutions of Type I with that on Type IV indicates that in both cases greatest growth was 1933J WYND — ORCHID SEEDLINGS AND NUTRITIONAL IONS 233 associated with the lowest proportioDs of the phosphate salt, irrespective of which ion it accompanied, and that the relative proportions of all other ions were of comparatively little effect. The fact that growth was inversely proportional to the concen- tration of the H 2 P0 4 ~ ion seems to be related significantly to the results of Gregory ('28). This author studied the effect of the different ions on growth by an ingenious mathematical treatment of the results published previously by Johnston ('24) concerning the growth of potato plants in three-salt solutions. In studying duplicate series, Gregory found that in each the fraction of the total growth produced by the H 2 POr ion was a negative quantity. K x SO H K*.SO t Fig. 5. Type IV, series 1, showing Fig. 6. Type IV, series 2, showing areas of high, medium, and low yield. areas of high, medium, and low yield. The cultures of high concentration The cultures of high concentration Ca(H,P0 4 ) 2 omitted (see text). Ca(HjPO«)» omitted. In general, he also found that the growth produced by other negative ions was much less than that produced by the positive ions. If we indicate the concentration of each ion in a "molar" sense and compare the composition of the best solution of Type I and Type IV in table vii, an interesting analogy is apparent. Despite the great apparent variation in composition of these two solutions, we see that in both the concentrations of the negative ions are of the same order of magnitude, while those of the positive vary seems therefore probable that growth 234 [Vol. 20 ANNALS OF THE MISSOURI BOTANICAL GARDEN being conditioned in both cases by the negative rather than the positive ions. TABLE VII CONCENTRATION OP IONS IN "MOLS" OP THE BEST SOLUTION OF TYPE I AND TYPE IV Type IV .0038 .0078 .0097 .0194 .0019 .0039 These results need not be interpreted as contradictory to those of Smith who found that better growth was obtained having larger amounts of ammonium sulphate added Knudson's solution B. Smith added nitrogen in the ammonium ion, while we used the It is entirely possible that orchid seedlings might react favorably to an increase of as the ammonium ion, while growth would the form of improve with This is indeed made very prob able by the work of Burgeff ('09), who found better growth with ammonium than with nitrate salts. We are at a loss to relate these results to those of La Garde ('29) who states, in regard to superiority of his solution L over that of Knudson's solution B, as follows The seedlings d darker in color and had progressed relatively further in opment. This effect might be ascribed to the larger dose dex of phosphorus and Experiment. which will be reported later, show that La Garde's solution I indeed a very favorable orchid medium, but it seems probable tl we may not ascribe this quality to the increase of phospho] and potassium per se. This study further emphasizes the suggestion made bv soi authors that thought effect Ca:Mg ratio is not of such importance as it , At least the limits of variation used in our not sufficiently great to produce any significant 1933] WYND — ORCHID SEEDLINGS AND NUTRITIONAL IONS 235 V. Summary Seeds of Cattleya Trianae Linden and Rchb. f. were germinated on three-salt solutions of Type I and Type IV, with the following results : — (1) The seedlings showed best development on media having low proportions of the total osmotic concentration due to the phosphate salt, irrespective of whether it was present as the potassium of the calcium compound. (2) The proportion of all other ions within the limits used appeared to have relatively little effect on germination and growth. (3) The negative ions appeared of greater importance in con- ditioning orchid seedling development than the positive ions. (4) The best growth in Type I solutions was found in culture Rl S2 having the composition: — KH 2 PO« 0025 M Ca(NO,) 2 0049 M MgSO* 0123 M (5) The best growth in Type IV solutions occurred in culture R5 SI containing: — K 2 SO< 0097 M Ca(H 2 P0 4 ) 2 0019 M Mg(NO a ) 2 0039 M VI. Acknowledgements The author wishes to thank Dr. George T. Moore, Director of the Missouri Botanical Garden, for furnishing seeds for this investigation; and also to express his appreciation to Elinor Al- berts Linder, formerly Orchidologist to the Missouri Botanical Garden, for helpful suggestions while employed at this institu- tion; and also to Dr. E. S. Reynolds, under whose supervision the work was carried out. 236 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 20 Bibliography Schreinemakers, F. A. H. ('93). Konzentrierung odor Verdiinnung einer Losung bei konstanter Temperatur. Zeit. Phys. Chem. 11: 81-85. 1893. Bancroft, A. D. ('02). Synthetic analysis of solid phases. Jour. Phys. Chem. 6: 178- . 1902. BurgefT, II. ('09). Die Wurzelpilze der Orchideen; ihre Kultur und ihr Leben in der Pflanze. Jena, 1909. Schreiner, O., and Skinner, J. J. (40). Ratio of phosphate, nitrate, and potaasium in absorption and growth. Bot. Gaz. 50: 1-30. 1910. Shive, J. W. ('15). A three salt nutrient solution for plants. Am. Jour. Bot. 2: 157-160. 1915. Wilson, J. K. ('15). Calcium hypochlorite as a seed sterilizer. Ibid. 420-427. 1915. McCall, A. G. ('16). Physiological balance of nutrient solution for plants in sand cultures. Soil Sci. 2: 207-253. 1916. e, J. W. solution. 1-63. 1917. , and Martin, W. H. ('18). A comparative study of salt requirements for young and for mature buckwheat plants in solution cultures. Jour. Agr. Res. 14: 151-175. 1918. *~~ -i y ('18a). A comparison of salt requirements for young and for mature buckwheat plants in water culture and sand culture. Am. Jour. Bot. 5: 186-191. 1918. ngstori, B. E., and Tottingham, W. E. ('18). A new three-salt nutrient solution for plant cultures. Ibid. 337-346. 1918. ■ , ('19). A plan for comparative research on the salt requirements of repre- sentative agricultural plants. Baltimore, 1919. Van Alstine, E. ('19). The relation of salt proportion to the growth of wheat in sand cultures. N. J. Agr. Exp. Sta. 40: 366-374. 1919. Martin, W. II., and Shive, J. W. ('20). A study of the salt requirements of the potato. N. J. Agr. Exp. Sta. Rept. 41: 409-412. 1920. Meier, II. F. A., and Halstead, C. E. ('21). Hydrogen-ion concentration relations in a three-salt solution. Soil Sci. 11: 325-352. 1921. Tottingham, W. E., and Rankin, E. J. ('22). Nutrient solutions for wheat. Am. Jour. Bot. 9: 270-276. 1922. Knudson, L. ('22). Non-symbiotic germination of orchid seeds. Bot. Gaz. 73: 1-25. 1922. Poole, R. F., and Pant, G. W. ('22). Further study of the relation of various fertilizers mixtures to the growth of celery in muck soil. N. J. Agr. Exp. Sta. Rept. 43: 395-399. 1922. Clement, E. ('24). Germination of Odontoglossum and other seed without fungal aid. Orchid Rev. 32: 233-239. 1924. , ('24a). The non-symbiotic germination of orchid seeds. Ibid. 359- 366. 1924. Ballion, M., and Ballion, G. ('24). The non-symbiotic germination of orchid seeds in Belgium. Ibid. 305-309. 1924. Johnston, E. S. ('24). Growth of potato plants in sand cultures treated with the "six types" of nutrient solutions. Md. A« r . Exp. Sta. Bull. 270: 53-86. 1924. Davidson, O. W. ('28). An application of the triangle system in determining a nutrient solution suitable for research with the peach in sand culture. Am. Soc. Hort. Sci., Proc. 25: 354-358. 1928. 1933] WYND ORCHID SEEDLINGS AND NUTRITIONAL IONS 237 Gregory, F. G. ('28). The differential effect of the ions of three-salt solutions on the growth of potato plants in sand cultures. Roy. Soc. Lond., Proc. B. 102: 311— 327. 1928. La Garde, R. ('29). Non-symbiotic germination of orchids. Ann. Mo. Bot. Gard. 16: 499-514. 1929. Quednow, K. G. ('30). Beitrage zur Frage der Aufnahme geloster Kohlenstoffver- bindungen durch Orchideen und andere Pflanzen. Bot. Archiv. 30 : 51-108. 1930. Addoms, Ruth M., and Mounce, F. C. ('31). Notes on the nutrient requirement and the histology of the cranberry (Vaccinium macrocarpon Ait.) with special reference to mycorrhiza. Plant Physiol. 6: 653-669. 1931. Smith, F. E. V. ('32). Raising orchid seedlings asymbiotically under tropical conditions. Gard. Chron. 91 : 9-11. 1932. Annals of the Missouri Botanical Garden Vol. 20 APRIL, 1933 No. 2 A REVISION OF THE NORTH AMERICAN SPECIES OF VERBENA 1 LILY M. PERRY Instructor in Botany, University of Georgia, Athens, Georgia Formerly Rufus J. Lackland Research Fellow in Botany in the Henry Shaw School of Botany of Washington University Introduction Verbena, the type-genus of the family Verbenaceae, is pre- eminently American. Owing to inherent variability and varia- tions caused by adaptation to local conditions of growth, the group is a rather complex one. Moreover, since the last comprehensive monograph of the genus was published so long ago that it contains only some twenty of the hundred or more described North American species and varieties, considerable material has been incorrectly named. It appeared, therefore, that a critical examination of the group might be helpful in untangling the existing confusion of identities. The following paper is an at- tempt to present the results of a study of the North American species, together with a brief review of the historical, morpho- logical, and phylogenetic aspects of the species under considera- tion. The materials studied are indicated in the text by abbreviations for the herbaria to which they belong: the Academy of Natural Sciences of Philadelphia (ANSP); Field Museum of Natural History (F); Gray Herbarium of Harvard University (G); the Royal Botanic Gardens, Kew (K); the Missouri Botanical 1 An investigation curried out at the Missouri Botanical Garden in the Graduate Laboratory of the Henry Shaw School of Botany of Washington University, and submitted as a thesis in partial fulfillment of the requirements for the degree of doctor of philosophy in the Henry Shaw School of Botany of Washington University. Issued July 10, 1933. Ann. Mo. Bot. Gard., Vol. 20, 1933. (239) [Vol. 20 240 ANNALS OF THE MISSOURI BOTANICAL GARDEN Garden (MBG); New York Botanical Garden (NY); Pomona College (P); United States National Herbarium (US); photo- graphs of types and authentic specimens have been received from the Museum of Natural History, Paris (Par.); the Herbarium of the Botanical Garden and Museum, Berlin- 1 lahlem (Bot. Mus. Berl.-Dahl.), and the Herbarium of the Botanical Garden, Madrid (Herb. Bot. Gard. Madrid). For the photographs and the use of the loans, as well as for the many courtesies received while visiting different herbaria, the writer is I ruly grateful and hereby acknowledges her indebtedness to all who have given assistance in this way. She expresses her hearty thanks and cordial appreciation to Dr. George T. Moore, Director of the Missouri Botanical Garden, for the use of the extensive privileges of the excellent library and the herbarium. She is also indebted to Miss Nell Horner, Librarian, for any suggestions in bibliographical matters. To Dr. J. M. Greenman, who has directed and supervised the work, giving ready aid and helpful suggestions at all times, the writer expresses her particular gratitude for the encouragement and personal interest shown throughout this study. m Paxonomic History The derivation of the word Verbena is obscure. The name has been handed down to us as representing certain herbs used in sacred rites by the ancients. Although apparently one of the well-known plants in botanical history, Verbena was described comprehensively for the first time by Tournefort, 1 whose descrip- tion formed the basis of the synopsis of the genus as accepted by Linnaeus. 2 In the { Species Plantaruni/ Linnaeus 3 records fourteen species of which eight have been removed since to other genera. The latter fact is sufficient to indicate that the Linnaean concept was too inclusive. It was held, however, until 1806, when Jussieu, 4 in an article on " Observationes sur la famille des plantes 1 Tournefort, Institutiones Rei Herbariae, 200, pi. 94- 1700. 1 Linnaeus, Gen. PL 12. 1754. 3 Linnaeus, Sp. PL 18-21. 1753. 4 Jussieu. Ann. Mus. Paris 7: 71-7:>. 1806. 1933] PERRY — NORTH AMERICAN SPECIES OF VERBENA 241 Verbenacees," pointed out that the genus was a composite one containing several units. Amongst these he showed that Verbena differed from the others in having four stamens and four nutlets. This definition was accepted by Endlicher 5 under the heading "verbenae verae," and it has continued to the present time. Although the genus is predominantly American, the earlier species were described largely from plants grown in various European gardens. This was unfortunate, since such types are often extinct or inaccessible or sometimes difficult to interpret on account of probable fusion or other modifications appearing in the species after some few years of cultivation. The first treatment of the North American species, as such, is that of Michaux. 6 He recognized eight species, all from the region of the middle and southeastern United States. From time to time new species were collected by pioneer explorers. Kunth 7 de- scribed eight from the Mexican region, Bentham 8 added three more, and Martens and Galeotti 9 reported eleven, of which six were new. The ablest and most inclusive work on the genus as a whole is that of Schauer. 10 In this he accepted seventy-one species and listed fourteen more as insufficiently known ; of these, twenty-seven are indigenous to North America. Appearing about the same time as Schauer's work is that of Walpers, 11 whose review might be considered as an assembling of the litera- ture of the genus rather than the results of a special monographic study; nevertheless, it is comprehensive and contains a few new species. Subsequently no publication of the North American species in toto has appeared, although the works of Gray 12 and Small 13 are very helpful in so far as regional floras are concerned. The last two publications contain the only keys to the group under consideration. The type-species of the genus is V. officinalis L. 5 Endlicher, Gen. PI. 633 (no. 3685). 1838. • Michaux, Fl. Bor.-Am. 2: 13-15. 1803. 7 HBK. Nov. Gen. et Sp. 2: 272-277. 1818. 8 Bentham, PI. Hartw. 21. 1839. « Martens & Galeotti, Bull. Acad. Brux. II 2 : 320-324. 1844. 10 Schauer in DC. Prodr. 1 1 : 535-556. 1847. "Walpers, Rep. 4: 13-33. 1844-48; 6: 686-687. 1846-47. 12 Gray, Syn. Fl. N. Am. 2 l : 335-338. 1878. "Small, Fl. Southeast. U. S. ed. 1, 1008-1011. 1903. [Vol. 20 242 ANNALS OF THE MISSOURI HOTANICAL GARDEN Gross Morphology Roots. — Some species are annual with a fibrous root system branching from a slender primary root; others are perennial with a larger tap-root. They maintain themselves from year to year by crowns from which clustered new shoots rise near the base of the old ones. A few species show a tendency to root at the nodes. Probably, for the most part, the main purpose of these secondary roots is to aid in obtaining nourishment for the plant, as instance, V. canadensis, a hardy plant with stems rooting only at the lower nodes. Nevertheless, in V. teucriifolia, they may be a possible means of propagation, as definite roots are found at more or less remote nodes; occasionally, both roots and fruiting spike are developed from the same node. In many cases, it is difficult to tell whether a species is an annual or only a short-lived perennial. The question of duration depends in some measure on climatic factors. Stems. — The stems are slender, terete, or, more commonly, quadrangular in cross-section with sharp or obtuse angles, and chiefly pubescent. Apparently all are herbaceous, the perennials dying down to the ground in winter and reappearing again in the spring. The shorter forms branch freely with decumbent- ascending or prostrate branches; the taller are usually erect with branches loosely ascending-spreading. All the lower branches are opposite, but the upper may be somewhat irregularly placed. Leaves. — In the majority of the species of the section Glandu- laria the leaves are deeply cleft, whereas those of the Vtrbenaca are in large part merely dentate or incised. As a whole they are membranaceous, varying greatly in size and somewhat in thick- ness. In a few species they are strictly sessile; in most, however, they are either subsessile or short-pet ioled and tend to be strongly veined beneath. All have more or less short, sheathing, and connate bases. In general the leaves are opposite, but at times they are subalternate or irregularly disposed on the upper part of the stem; they are rarely ternate, and this arrangement appears to be an individual expression rather than a specific or even a varietal character. Pubescence. — The trichomes are simple and unicellular, varying in quantity, length, rigidity, coarseness, and direction. None 19331 PERRY — NORTH AMERICAN SPECIES OF VERBENA 243 of the pubescence is truly hispid, but in some species the hairs are slightly stiff er than in others; this indument, as exemplified in the calyx of V. bipinnatifida y is designated as "hispid-hirsute." In V. prostrata and V. Gooddingii the pubescence is character- istically soft- villous, but in the majority of species the general tendency is to approach a hispidulous-hirsute to strigillose condition. In the former instance, the trichomes, especially along the stem and on the inflorescence, are of different lengths and more or less spreading; whereas in the latter, as in V. tarn- pensis, they are short, approximately equal, and appressed. Often the hairs of the upper surface of the leaf spring from minute bulbous bases; these, according to Solereder, 14 are calcified or silicified hairs, and the trichomes with somewhat pustulate bases, in V. Carolina and occasionally in other species, are cystolith- hairs. In any case, Solereder did not find cystoliths dissociated from the hairs. A viscid condition is rarely present, but glandular hairs are widely distributed throughout the genus. Inflorescence. — The inflorescence is terminal at the apices of stems or branches. The flowers are subtended by bracts and are borne in solitary or somewhat panicled or cymosely arranged spikes. Although these vary considerably, it is fairly easy to determine the section to which a plant belongs by the type of its spike. In the majority of the indigenous North American species of the Verbenaca, the spikes are elongated and tend to be irregularly placed, as in Verbena Carolina, so that they appear more or less panicled. In V. sphaerocarpa and V. litoralis, however, they are commonly arranged in simple or compound cymes. A spike may be either very slender with the flowers somewhat remote, or dense with imbricated flowers. In general, the elongated and comparatively narrow spike with small and inconspicuous flowers is a distinctive feature of the Verbenaca native to our continent. On the other hand, the spikes characteristic of the section Glandularia are shorter and relatively broad with larger and more showy flowers. In the earlier stages of anthesis these are in fascicle-like clusters; later, when the rhachis elongates, the in- florescence becomes spicate. "Solereder, Syst. Anat. Dicot. ed. 2, 1: 631-633. 1908. 244 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 20 Calyx.— The calyx is tubular, 5-lobed or subtruncate, 5- nerved and persistent. The nerves protrude somewhat beyond the margin of the lobes, forming calyx-teeth; these vary more or less specifically and occasionally intraspecifically. The nerves are unequal in length and the posterior one, along which the calyx ruptures to release the schizocarp, is always the shortest. In the Glandularia, the calyx is at least twice as long as the schizocarp and at maturity the teeth and lobes are more or less contorted. In contrast, the calyx of the Verbcnaca is generally shorter and may be either connivent, concealing the schizocarp as in V. scabra, or, as in such species as V. urticifolia and V. officinalis, open, disclosing the apices of the nutlets. With the developing ovary, the calyx enlarges a little at the base; usually it is somewhat angled, but in V. tumidula it is round at the base and tends to be inflated. Although only relative in value, the pubescence of the calyx is probably of more worth in specific determination than the other previously mentioned characters. Corolla. — The corolla is salverform and various in color. The tube is either straight or incurved, as long as or much longer than the calyx, and uniform or slightly enlarged at the throat. The degree of 1 lie exsertion of the corolla- tube is apparently a conveni- ent and obvious character, but one to be handled with care; first, it is somewhat variable, and second, when the corolla falls after an thesis, it does not drop at once but may be some little time working its way out of the calyx-tube; hence, the length of the exserted portion appears much greater than it really is. The limb is lobed and may be either conspicuous (6-12 mm. broad) or inconspicuous (2 mm. broad). The shape and the direction of the lobes may be of value in field identification but are not at all satisfactory in work on herbarium material. The throat inside is bearded with straight minutely roughened retrorse hairs and at the orifice with moniliform ones; outside and beyond the calyx it is either glabrous or finely pubescent. The inner hairiness is more prominent in the section Glandularia. Stamens. — The stamens are didynamous, often very short- stalked, and in the North American species, inserted on the upper middle half of the corolla-tube, but not exserled. In V. bipin- natifida and all the related species of its section, the anterior pair 1933] PERRY — NORTH AMERICAN SPECIES OF VERBENA 245 is very near the orifice of the throat and usually bears a gland- like appendage on the connective. The stamens are too much alike to be useful in specific differentiation. Pistil. — The ovary is superior, bicarpellary, and entire or very shallowly lobed at the apex. The style varies in length with the length of the corolla-tube, bearing the mature stigma approxi- mately in the region of the lower anthers. The typical form is slender and bilobed, the posterior lobe being sterile and laterally compressed, the anterior stigmatic and broad. The relation of these lobes to each other is reasonably constant throughout each section of the genus. In V. scabra, however, the broad stigmatic surface appears to be subtended on either side by a sterile style- lobe. Another interesting variation of the style is produced by V. quadrangulata ; here, the base is enlarged or thickened into a somewhat hemispherical-angulate stylopodium and persists as an integral portion of the nutlets. Fruit — The fruit of Verbena is a schizocarp enveloped by the persistent calyx. As previously stated, the calyx splits along the posterior nerve to release the schizocarp attached to a more or less round-quadrangular gynobase. For the most part, four nutlets or pyrenae develop, two from each biovulate carpel. Generally the nutlets are either subcylindric or subtrigonous, but in V. quadrangulata they are beaked cylindric, and in V. tumidula subovoid. The dorsal surface is more or less convex with the lateral surfaces scarcely definite, rather appearing as continuations of the dorsal portion and adjoining the commis- sural face or faces. This is not always the case, since in V. tumidula the lower part of the nutlet seems inflated and the lateral is somewhat ventricose and smooth. The surface of the nutlet varies from essentially smooth, as in V. Carolina and its to striate or sulcate, and is oftenest raised-reticulate or ate-scrobiculate on the upper part. Occasionally, the character or scope of these depressions is of specific value, as in V. gracilis; but usually a certain type of nutlet is characteristic of a group of species rather than a single one. The commissure may be more or less muricate or practically smooth. In the subcylindric pyrenae of the section Glandularis, the commissural face is commonlv convex and does not extend allies, 240 ANNALS OF THE MISSOURI HOTANICAL GARDEN [Vol. 20 to the tip of the nutlet; whereas in the trigonous nutlets of the Verbcnuca, the commissure ordinarily has two faces meeting at right angles, and is as long as the nutlet. The gynobase varies with the size and shape of the pyrenae which it bears. In V. Carolina and V. Ehrenbergiana, it is nearly flat and almost orbicular, but in species with larger nutlets, as in V. simplex and V. neomexicana, it is slightly deeper and in large part somewhat quadrangular. It shows its greatest develop- ment in the section Glandularia where the pyrenae are largest and often somewhat broadened at the base. Perhaps the gyno- base should be used as a character complementary to that of the nutlets, but its definite value is rather elusive. The more important morphological characters, which have been used in the delineation of the genus and its sections, are illustrated with a legend (pi. 13). Hylrrids. — It is a well-accepted fact that the cultivated races of Verbena have arisen through t he more or less fusing of species attractive for horticultural purposes. Granting this, without field observation and knowledge gained through experimental work, the writer does not propose to discuss the question of hybridization in Verbena. Nevertheless, it is necessary to point out that there is a considerable number of specimens amongst the collections of this genus from the Middle States, which show variation in several directions or a combination of the characters of two or more species; it is the prevailing opinion that these plants are hvbrids. Relationships and Distribution The genus Verbena is a member of the tribe Verbeneae accord- ing to Engler and Gilg, 15 or of the Euverbeneae according to Briquet in Engler and Prantl." It is distinguished from its immediate relatives by the four dry and hardened nutlets. In this paper fifty-one species are recognized, four naturalized and forty-seven indigenous. With the exception of a few border-line species, it is relatively easy to separate the genus into two fairly distinct sections in which the majority of the members are so 15 Engler A Gilg, Syllab. dor Pflanzonf. eds. «.) A 10, 339. 1924. " : Engler & Prantl. Nut. Pflansenf. 4 s *: 14<>. 1895. 1933] PERRY — NORTH AMERICAN SPECIES OF VERBENA 247 different that it would seem as if they had followed divergent trends for a long period of time. In attempting to present any genus in a natural order, it is highly desirable to know what species are primitive. However, without an intimate knowledge of the more diversified South American species, it seems prefer- able to state the evidence derived from the study of the North American group and leave the conclusions vague rather than more or less arbitrary. The section Verbenaca is probably the older of the two. It has the wider distribution, ranging from Canada to Panama as well as from the east to the west coast of United States and Mex- ico. Also, it is larger, containing thirty-two species. A number of these are annual, the remainder are perennial, but it is often hard to distinguish the members by this criterion. Although admittedly varying and apparently intergrading, the species as a whole are more stable and more readily recognized than those of the section Glandular ia . The flowers are much smaller and for the most part are produced in greater abundance; the nutlets also are abundant and in some cases minute. The combination of these characters leads one to conclude that this section is the more primitive one. Within the section, some species appear to be more closely related than others. V. officinalis, V. menthaef olia , V. Halei, and V. riparia form a natural group possessing the same general habit, type of pubescence and of nutlets. These are species of more or less adjacent regions except V. officinalis , which has been introduced from Europe. Again, V. Ehrenbergiana, V. Carolina, V. longifolia, and V. recta are very closely related, with V. urticifolia and V. scabra only a little farther removed. The last five species differ from the first mentioned in their more or less upright habit and less divided leaves, also in their smaller and somewhat smoother nutlets. It would appear as if all had arisen very close to the ancestral form and diverged only a little in the process of adaptation to environment. V. hastata and V. simplex are quite distinctive in habit, but the abundance of intermediate forms occurring between these and the two species, V. stricta and V. bracteata, would seem to indicate a close relationship amongst these units. V. Orcuttiana, although 248 ANNALS OF THE MISSOURI UOTANICAL GARDEN [Vol. 20 somewhat isolated, appears to have its nearest affinity in V. simplex. A third closely related group contains V. canescens, V. neo- mexicana, V. plicata, and V. gracilis; the common characters here are those of leaf-incision, pubescence, and general features of inflorescence. V. percnnis resembles this group in all respects except in its quite aberrant pubescence of short antrorse and hispidulous trichomes; in this character it simulates more closely the first-mentioned group. The following are more or less isolated species of limited distribution and with fairly static characters: V. xutha, V. prostrata, and V. robusta. V. cornea is probably a relic of some ancient form; the distinctive character of the schizocarp gives no clue to its affinities, but rather emphasizes the anomaly of the species. The section Glandularia seems to be the more nascent. The species are perennial, developing from a crown, and have a more or less sprawling to upright habit. The characters are highly variable and some species are difficult to distinguish, apparently passing into each other. Particularly is this true of V. ambrosi- folia, V. Wrightii, and V. ciliata. Some of the specimens from northern Mexico have been most perplexing; while it is perfectly evident that they belong to this group, none of them is typical. V. bipinnatifida is a closely related and wide-ranging species; by comparison, its characters are fairly clear-cut. The remaining species of the section are easily associated in groups or are anomalous. V. canadensis, V. mar i lima, and V. tampensis are very much alike either in habit or in floral structure. V. tam- pensis, however, is one of the species of this group where the glands may or may not be present on the anthers, and may represent the development of some transitional stage between the two sections. V. elegans of Mexico is very closely parallel to V. canadensis of the southeastern and central United States. V. delticola seems to belong in this association also; yet it is interest- ing to note that its fruit shows a tendency to develop a beak, and this may be significant of some little affinity with V. quad- rangulala, a species so aberrant that it has been treated as a separate genus. V. Gooddingii, V. lumidula, and V. pumila 1933] PERRY — NORTH AMERICAN SPECIES OF VERBENA 249 might very well be related to any of the previously mentioned species of the section. V. setacea resembles V. Gooddingii var. nepetifolia in habit, but it has such long calyx-teeth that one is a little hesitant about expressing any ideas of its relationship. V. teucriifolia and V. ciliata appear to intergrade in Mexico. The former, as well as V. pumila, shows a tendency toward reduced corollas and perhaps parallel development. V. lilacina and V. amoena are without close relatives. Considering the genus as a whole, some species are in a state of flux showing many atypical forms and covering fairly large geographic areas; others are just as widespread and yet are characterized by more static characters; and again, others are endemic showing a very limited distribution and appearing as outlying members of the group. The species probably vary from youth to age in the order mentioned. As previously stated, the genus Verbena is fairly widely distributed in North America; its range extends from southern Canada to Panama and from the east coast to the west both in United States and in Mexico. It is found also in the West Indies and on islands in the Pacific off the coasts of California and Mexico. It occurs in great abundance in Texas, New Mexico, and Arizona, south along the Sierra Madre and throughout the eastern part of Mexico. It is also fairly well distributed in southern Mexico, but only a few species are reported from Central America. The region of greatest specific concentration for North America is in the state of Texas, where twenty out of the forty-seven indigenous North American species are rep- resented. Amongst these species are members of both sections, some with very definitely clear-cut characters and more or less limited distribution, others with fluctuating characters and more widely distributed. By way of contrast, let us consider the outlying members of the genus which have stable characters and are isolated or in- habit very limited areas — V. lilacina, V. sphaerocarpa, V. Or- cuttiana, V. setacea, V. macrodonta, V. amoena, and V. subuligera; or again species such as V. carnea and V. quadrangulata, which are singular but have a larger geographic range. All these appear to be relics of a more ancient distribution. Undoubtedly 250 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 20 V. lilacina and V. sphaerocarpa, endemics of Cedros and Socorro Islands respectively, developed as a result of geographic isolation. The same may be true of the Lower Calif ornian entities, V. macrodonta, V. selacea, and V. Orcutliana. Perhaps V, amoena from Lecheria, a locality some little distance north of the city of Mexico, and V. subuligera, from t he Sierra Madre west of Durango, owe bolh their restriction and their differentiation to the presence of mountainous barriers. However, the same conclusion is hardly valid for either V. cornea, ranging from North Carolina to Florida and west to Texas, or V. quadrangulata, of southern Texas and Tamaulipas; yet they are probably relic endemics which have survived in favoring areas. If this be true, here is another bit of interesting evidence supporting the idea that the Verbcnaca is the older of the two sections. Of the nine species enumerated above as being of more ancient derivation, four, V. setacea, V. lilacina, V. amoena, and V. quadrangulata, have been placed in the section Glandularia; yet, in the flowers examined, the anthers are definitely unappendaged. To be sure, this disposition of the species is not in accord with the previous classifications and may not be acceptable; nevertheless, it seems logical to believe there must be transitional forms in the develop- ment and evolution of species; and in this case, the major portion of the characters of the species in question are those of the section Glandularia. Two other species which have developed a little farther along this same line are V. tampensis and V. pumila; gland-like appendages may or may not be present on the anthers; if present, they are usually very small. It may be of interest to note that, in agreement with Jordan's law of distribution, the species associated in groups usually occupy adjacent regions. Now as to origin.— It is possible that the progenitors of Verbena were very widespread at the beginning of the Pleistocene and with the glacial advance migrated southward; later, when more favorable climatic conditions developed they spread northward, establishing themselves in the present regions of distribution. Another possibility is that the centre of distribution is in South America and migration has been northward. However this may be, V. litoralis is the only known species indigenous to both 1933] PERRY — NORTH AMERICAN SPECIES OF VERBENA 251 countries and does not seem to be very closely related to the remaining North American species. Without a detailed in- vestigation of the South American group, it seems preferable to reserve assumptions in the matter. Economic Value Verbena is of very little value economically. It was introduced into cultivation in European gardens relatively early in the his- tory of botanical explorations. Although not so popular as it was once, on account of its vigor and hardiness, it is still used in many gardens to impart bright colors to the landscape. Taxonomy Verbena [Tourn.] Linn. Gen. PL 12. 1754. Obletia Rozier, Jour. Phys. 1 : 367, pi. 2. 1773. Glandularia Gmel. Syst. Veg. 920. 1791. Billardiera Moench, Meth. 369. 1794. Shuttleworthia Meisn. PL Vase. Gen. 290. 1839, and Comm. 198. 1839. Uwarowia Bunge, Acad. St. Petersb. Bull. Sci. 7: 278. 1840. Calyx tubular, 5-toothed, 5-nerved, at maturity often slightly enlarged at the base, persistent. Corolla minute to conspicuous, hypocraterimorphous, subbilabiate, 5-lobed; tube erect or in- curved, uniform or slightly enlarged above, equalling the calyx- lobes or surpassing them; lobes imbricate in the bud, broadly oblong, obtuse or retuse, unequal. Stamens 4, didynamous, included; anthers ovate, unappendaged or the connective of the upper pair expanding extrorsely into a clavate and gland-like appendage; filaments usually short, attached above the base of the corolla-tube. Ovary superior, entire at apex or very shortly 4-lobed, bicarpellary, 4-loculed; ovules in pairs, one to each locule, anatropous; style terminal, mostly bilobed, the anterior lobe stigmatose and the posterior usually sterile. Schizocarp included in the persistent calyx, dry, hardened or brittle, separat- ing into 4 homomorphous nutlets. Seed erect; embryo straight; endosperm none. — Chiefly American. Herbs or shrubs, erect or decumbent to prostrate, pubescent or occasionally glabrous. Leaves opposite, rarely ternate or verticillate, or the upper 252 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 20 alternate (irregular), variously dentate, incised or dissected, not often entire. Spikes terminal, rarely axillary, peduncled or sessile, densely flowered or elongate and slender with flowers remote, solitary, cyinose or paniculate. Flowers hermaphroditic, zygoniorplfir, sessile and bracteate. Section 1. Verbenaca Schauer 1. Verbenaca Schauer in DC. Prodr. 11: 536. 1847. Sterile style-lobe adjacent to the stigmatic surface or sub- lateral but usually not protruding beyond it; ovary entire at the apex with the style attached at the distal end, or, if at all lobed, style inserted in the indefinite depression between the lobes; schizocarp commonly not constricted along the lines of cleavage; anthers unappendaged. Annuals or herbaceous perennials with prostrate, decumbent-ascending or erect stems. Flowers mostly not showy. Calyx rarely more than twice as long as the schizo- carp and not contorted beyond it. Species 1-32. Ser. I . PACHYBTACHYAE Schauer. I leads or spikes, at least in anthcsis, crowded and short, not at any time greatly elongated or open, generally disposed in compound cymes. — Chiefly introduced South American species. Key to the Species A. Leaves scmiamplexicaul and subcordate. B. Inflorescence glandular; bracts conspicuously longer than the calyx; corolla-tube 2-3 times as long as the calyx /. V. rigida B. Inflorescence not glandular; bracts barely equalling or only slightly ex- ceeding the calyx; corolla-tube scarcely twice as long as the calyx. . . . 2. V. bonariensis A. Leaves not semiamplexicaul nor subcordate, tapering into a cuneate- attenuate subsessile or petiolar base. C. Spikes short (3-5 mm. long), very dense and appressed-pubescent. D. Fruiting-calyx ascending; schizocarp longer than broad, raised- reticulate above, striate toward the base 3. V. brasiliensi HIS I). Fruiting-calyx spreading; schizocarp as broad as long, practically smooth J i . V. sphacrocarpa C. Spikes 5-10 mm. long, dense at the apex, somewhat open below and finely strigillose 5. V. litoralis 1. V. rigida Spreng. in L. Syst., Cur. Post. 4 2 : 230. 1827. V. venom Gill. & Hook. Hot. Misc. 1: 167. 1830. 1933] PERRY— NORTH AMERICAN SPECIES OF VERBENA 253 V. bonariensis var. rigida 0. Kuntze, Rev. Gen. 3 2 : 255. 1898; Briq. Ann. Conserv. & Jard. Bot. Geneve 7-8: 291. 1904. Stems 2-6 dm. tall, sharply 4-angled in cross-section, scabrous- pubescent; leaves oblong to oblong-lanceolate or narrowly obovate, sessile, more or less semiamplexicaul, subcordate, sharply and coarsely serrate, midrib and veins impressed above but prominently reticulated beneath, scabrous and more or less hispidulous on both surfaces, margins of older leaves somewhat revolute; spikes usually short, dense and cylindrical, disposed in subternate cymes with the laterals pedunculate; bracts lanceolate, acuminate-subulate, usually closely imbricated and longer than the calyx, glandular-pubescent, ciliate; calyx 4 mm. long, glan- dular-pubescent, lobes acute with short mucronate-subulate tips; corolla-tube twice (-thrice) as long as the calyx, pubescent without; corolla-limb 5-7 mm. broad; stamens inserted on the lower middle of the corolla; nutlets trigonous, slightly broader at the base than at the apex, scarcely 2 mm. long, raised-reticulate on the upper half, striate toward the base; commissural faces muricately scabrous. West southern United States. Specimens examined: Randolph (G). Georgia: uncultivated ground, in perennial spreading patches, 26 May 1928, Reade (NY); vicinity of Thomson, McDuffie Co., 1 Sept. 1907, Bartlett 1122 (P). Alabama: Mobile, Mohr (US); Mobile, June 1919, Graves 525 (MBG, US). Louisiana: Mandeville, 15 Aug. 1912, Pennell 4204 (NY); Covington, Aug. 1919, Arsbne (US); Catalpa, 24 Aug. 1912, Pennell 4332 (NY); Baton Rouge, 28 March 1904, Billings 49 (G); roadsides, Plaquemines Co., July 1883, Langlois (NY). Texas: Orange, 1914, Young 662 (P, US); Huntsville, 24 May 1917, E. J. Palmer 12038 (MBG); Houston, 1872, Hall 433 (G, MBG, NY, P, US); Houston, 1913, Fisher 460, 625 (US); Houston, 25 Sept. 1915, Thurow (US); Houston, 22 May 1917, E. J. Palmer 12001 (MBG); 16 km. southeast of Houston, 12 April 1925, Small & Wherry 11813 (NY). Mexico: Vera Cruz: Zacuapan, May 1913, Purpus 6413 (F, G, MBG, NY, US). West Indies: Bermuda: St. George's, 13 July 1905, Moore 2984 (G); near Devil's Hole, 13 June 1905, Harshberger (ANSP, G, MBG); near Hamilton, 31 Aug.-21 Sept. 1905, Brown & Button 153 (ANSP); Tucker's Town, 3 May 1912, Collins 45 (G); Tucker's Town, 8 Aug. 1913, Collins 269 (G, NY); fields, Montrose, Sept. 1913, Brown, Brilton & Wortley 1645 (ANSP, NY); St. David's Island, 22 May-2 June 1919, Brown 693 (ANSP, NY). 254 ANNALS OF THE MISSOURI [Vol. 20 Cuba: near Farallones, Oriente, 5 Aug. 1913, Leon 8.910 (NY). Jamaica: road to Salt Hill, St. Andrew, 6 May 191. r >, Harris 11969 (G, MBG, NY); Blue Mountains, St. Andrew, 20 March 1916, l'crkins 102/, (G); near Fairfield, 3-7 Sept. 1908, Bntton 3176 (NY). Guadeloupe without locality, 1893, Duss 8+70 (NY). Maktiniqi i ■:: without locality, Sept. 1901, Duss 4697 (NY). This South American species is commonly cultivated. It has escaped and established itself in various places. Kuntze and Briquet have regarded it as a variety of V. bonarien sis , but lacking a considerable amount of material for examination, the writer prefers, at present, to maintain it as a sDecific entitv. 1827 2. V. bonariensis L. Sp. PI. 20. 1753. V. elongate Salisb. Prodr. 71. 17%. tV. quadrangular™ Vellozo, Fl. Flum. 1(>. 1825; 1 :pl 89. St (mi is 1 iii. more or less in height, 4-angled in cross-section, somewhat scabrous-pubescent; leaves decussately opposite, lance- olate, subauriculate, semiamplexicaul, acutely serrate, entire toward the base, rugose and hirtellous above, spreading-pubescent beneath, particularly along the prominently reticulated veins; spikes compact, mostly short, commonly sessile and crowded in dense fasciculate cymes; bracts lanceolate-acuminate, barely equalling or slightly exceeding the calyx, pubescent; calyx 3 mm. long, pubescent, lobes acute with short subulate tips: corolla-tube limb the length of the calyx, pubescent slightly raised-reticulate at apex; commissural faces scarcely reaching the tip of the nutlet, muricate-scabrous. Distribution: native of South America; introduced into wmthern United States and the West Indies. Specimens examined: South CAROLINA: roadsides near Charleston, Curtiss 1963 in part (F, G, MBG, NY); Charleston Neck, 28 Sept. 1853, Gibbcs (NY, US); Charleston, 30 April 1912, 7 Robinson 12 (MBG, NY). 148 GEORGIA: along canal bank, Augusta, Cuthbcrt 267, S5S (NY). Louisiana: vicinity of Covington, 1920, Arsene 118.59, 12634 (F, US). Aukansab: roadsides near Forrest City, 17 Oct. 1925, #. J. 1 'aimer 29303 (G.MBG, NY). Texah: Huntington, 15 Sept. 1923, Thorp 2558 (US). West Bermuda: waste ground, 29 March-3 May 1909, Marble 737 (ANSP, NY); Pern- 1933] PERRY — NORTH AMERICAN SPECIES OF VERBENA 255 broke Swamp, 21 June 1905, Harshberger (ANSP, G, MBG, NY); roadside, Pem- broke, 31 Aug.-20 Sept. 1905, Brown & Britton 98 (ANSP, G, NY, P); Hamilton, 12 July 1905, Moore 2946 (G); Hamilton, 16 Jan. 1912, Robinson 94 (G); Tucker's Town, Collins 53 (G), 266 (G, NY). Jamaica: Blue Mountains, 14 Jan. 1890, Hitchcock (MBG); Blue Mountains, Mt. Hybla, 5 Apr. 1916, Perkins 102 (G); St. Helen's Gap, 7 April 1909, Taylor 4244 (NY); St. Helen's Gap, St. Andrew, 4 March 1920, Maxon & Killip 569 (G, NY); Cinchona, 26 July 1903, Nichols 168 (G, MBG, NY); Cinchona, 22 Dec. 1905, Harris 9132 (NY); Arntully, 25 Aug. 1927, Orcutt 2736 (G, MBG). This is another introduced South American species evidently related to V. rigida, but it is a larger and coarser plant with smaller flowers as well as less harsh and somewhat viscid pubes- cence. 3. V. brasiliensis Vellozo, Fl. Flum. 17. 1825; 1 : pi. 40. 1827. V. litoralis var. brasiliensis Briq. Ann. Conserv. & Jard. Bot. Geneve 7-8: 292. 1904. Stems 1 m. more or less in height, 4-angled in cross-section, practically glabrous below, somewhat scabrous-pubescent above, slightly contracted at the nodes; leaves decussate, lanceolate, tapering into a cuneate-attenuate, subsessile or petiolar base, sharply or incised serrate, strigillose and somewhat pustulate above with veins impressed, sparsely pubescent beneath; spikes compact, mostly short and strict, usually sessile in open cymes; bracts scarcely as long as the calyx, lanceolate, subulate at apex, ciliate; calyx approximately 3 mm. long, somewhat appressed- pubescent, lobes acute with short subulate tips; corolla-tube a little longer than the calyx, pubescent without; corolla-limb inconspicuous; nutlets trigonous, about 2 mm. long, strongly striate, raised-reticulate at apex; commissural faces scarcely reaching the tip of the nutlet, muricate-scabrous. Distribution: South America; introduced into southern United States. Specimens examined: North Carolina: ballast dumps at Wilmington, 2 July 1897, Small 5707 (G, MBG, NY); dry sandy soil, open woods about 3 km. south of Wilmington, 25 July 1922, Randolph 1004 (G). Georgia: savanna, 27 May 1927, Korthoff (US); Brunswick, 12 May 1930, Mol- denke 1184 (MBG, NY). Florida: waste place, Chipley, 24 May 1930, Blanton 6598 (MBG, US); waste ground, Pensacola, 24 July 1899, Curtiss 6490 (G, MBG, NY, US); ballast ground, Pensacola, 30 April 1903, Tracy 8706 (F, G, MBG, NY, US). Alabama: ballast ground, estuary of Mobile River, 4 July 1893, Mohr (US); Port 256 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 20 Eads, 22 Aug. 1900, Lloyd & Tracy 20 (F, G, MBG, NY, US); north bank, Bayou Terrebonne, llouma, 29 May 1914, Wurzloxv (NY). A species very similar in habit to V. litoralis but readily distinguished by the difference in inflorescence. The spikes of V. brasilicnsis are short, compact, sessile, and regularly arranged in open cymes; whereas those of V. litoralis are longer, compact or somewhat elongate, peduncled, and arranged in more or less paniculate cymes. 4. V. sphaerocarpa Perry, 17 n. sp. Steins erect, square in cross-section, branched above, glabrous or sparsely scabrous-pubescent; leaves oblong, tapering at the base into a short petiole, 3-()(-10) cm. long, acute, sharply serrate, the upper often entire, scabrous-pubescent above with trichomes minutely bulbous at the bases, more or less short- strigillose on both surfaces, prominently veined beneath; spikes short and dense, subsessile or short-pedunculate, cymosely arranged ; bracts ovate-lanceolate, shorter than the calyx, acute, pubescent; calyx scarcely 2 mm. long, connivent over the schizo- carp, appressed-pubescent, teeth very short, acutish; corolla- tube protruding slightly beyond the calyx; corolla-limb about 1.5-2 mm. broad; schizocarp 1 mm. long, approximately 1 mm. in diameter; nutlets very faintly striate or essentially smooth; commissural faces muriculately scabrous. Distribution: known only from Socorro Island. Specimens examined: Mexico: Colima: Socorro Island: March June 1897, Anthony 380 (G, MBG type, US); 27 May-3 July 1903, Barkelew 231 (G, MBG, P, US); 9 May 1925, Solis 70 (US). This endemic from Socorro Island is most nearly related to V. 17 V. sphaerocarpa Perry, sp. nov., annua vel perennis (basi ignota); caule erecto quadrangulari glal>ro vel sparse scabro-pubescente; foliis oblongis basi in brevem petioluin attenuate 3-6(-10) cm. Iongis aculis, infimis argute serratis, superioribus saepe integris, omnibus scabro-pubescentibus supra subtusque plus minusve breviter strigosis et rcticulatis; spicis brcvibus et compactis subsessilibus vel breviter pe- dunculitis in cymas dispositis; bracteis ovato-lanceolatis acutis pubescentibus calyce brevioribus; calyce vix 2 mm. longo connivente adpresso-pubescente; calycis dentihus brevissimis acutiusculis; corollae tubo paulo exscrto; corollae limbo eirciter 1.5-2 mm. lato; schizoearpio 1 mm. alto eirciter 1 mm. Iato; coccis obsolete striato- reticulutis. — Collected on Socorro Island, Mexico, March June 1897, Anthony 380 (MBG), TYPE. 1933] PERRY — NORTH AMERICAN SPECIES OF VERBENA 257 litoralis and has a similar habit, but is easily separated on the distinctive characters of the inflorescence. The spikes are shorter and denser, the flowers are smaller and so crowded that the lower ones appear to be inserted at right angles to the rhachis of the spike; moreover, the schizocarp is fully as broad as or even broader than long, an unusual trait not found elsewhere in the North American species of Verbena. 5. V. litoralis HBK. Nov. Gen. et Sp. 2: 276, pi. 137. 1818. V. caracasana HBK. I. c. 275. V. bonariensis var. littoralis Hook. Bot. Misc. 1 : 166. 1830. V. affinis Mart. & Gal. Bull. Acad. Brux. II 2 : 322. 1844. V. littoralis a pycnostachya Schauer in DC. Prodr. 11: 542. 1847, excluding V. brasiliensis Veil. Fl. Flum. 1 : pi. 40. V. littoralis /3 leptostachya Schauer in DC. Prodr. I. c. V. Hanseni Greene, Pittonia 3 : 308. 1898. V. litoralis var. caracasana Briq. Ann. Conserv. & Jard. Bot. Geneve 7-8: 292. 1904. Stems approaching 1 m. in height, square in cross-section, somewhat fastigiately branched above, glabrous or sparsely strigillose, slightly contracted at the nodes; leaves lanceolate or oblong, tapering into a very short petiole or a subsessile base, 3-10 cm. long, decussate, more or less sharply and coarsely serrate, sparsely strigillose on both surfaces, scabrous and some- what rugose above, prominently veined beneath ; spikes terminal, fairly dense, pedunculate, cymosely arranged or tending to be panicled, often elongate; bracts ovate-lanceolate, acuminate, sub- equalling or somewhat shorter than the calyx, glabrate; calyx 2-2.5 mm. long, finely strigillose, subtruncate, teeth minute, subulate ; corolla-tube variable in length, always somewhat longer than the calyx; corolla-limb inconspicuous, 2.5-3 mm. broad; nutlets trigonous, hardly 2 mm. long, striate, somewhat reticulate at the apex; commissural faces about as long as the nutlet, muricate-scabrous . Distribution: Mexico, Central and South America; introduced into the United States. Specimens examined: Louisiana: north bank, Bayou Black, Houma, 5 and 9 May 1914, Wurzlow (NY). California: Clinton, Amador Co., June 1896, Hansen 2025 (MBG). [Voi-. a<) 258 ANNALS OF THE MISSOURI HOTANICAL GARDEN Mkxico: San Luis Potosi: on mountains around the city, San Luis Potosi, 1876, Schaffner 718 (G); Alvarez, 28 8ept.-3 Oct. 1902, Palmer HI in part (G). Nayahit: Tepic, 5 Jan.-6 Feb. L882, Palmer 9019 (l T S), 2014a (G, NY). Jalisco: fields, Guadalajara, 1 1 Aug. 1902, Pringle 11099 (F, G, MBG, NY, US). Hidalgo: near Ixmiq u i 1 pan , 1905, Rose, Pointer & Row9077, 9151 (US). Vbka Cru«: Orizaba, 27 July 1891, Scalon 27 (F, G, NY, US); Iluatusco, April 1857, Mohr (US); Fortin, Feb. 1883, Kerber 811 (US); Santa Rosa, 13 Aug. 1926, Fisher 168 (US). Tabasco: Mayito, 10 April 1889, Roviroaa (ANSP, NY, US). Pueula: Pahuatlan, 14 June 1913, Salatar (US); near Puebla, 20 Oct. 1908, Arsene. (US). MlCHOACAN: low valley, Zinapeeuaro, 2 May 1849, Gregg 764 (MBG); Quinceo, 11 Nov. 1909, Arsene (US); Morelia, Aug.-April 1840, OaleoUi 781 (K), type col- lection of V. affinis; Morelia, Coronilla, 8 Aug. 1909, Arsbne (US). Oaxaca: wet meadows, Sierra de San Felipe, 11 Sept. 1894, Pringle 4877 (ANSP, G, MBG, NY, P, US); Sierra de San Felipe, ti ( »et. 1894, C. L. Smith 222 (MBG, US); San Jacinto, 25 Sept. 1895, L. C. Smith 79 ', (G). Centual Amekica: Guatemala: uncultivated places, Oct. 1885, Bernoulli 128 (NY); without locality, 1892, Heyde, 610 (US); waste places near railway bridge, 5 June 1909, Deam 6180 (G, US); vicinity of Los Amates, Dept. Izabal, 24 May 1922, Standley 24404 (US); Coban, Dept Alta Verapaz, May 1886, von Twrckheirn 904 (ANSP, F, G, NY, US); near the Pinca Sepacuite, Dept. Alta Verapaz, 25 March 1902, Cook & Griggs 148 (US); Canjuti, Dept. San Marcs, 1 Sept. 1922, Solas 82 (US); Guatemala, 1923, Ruano 862, 408 (US); Guatemala Gity, 1912, Aguirre 4 (VH); near Guatemala, July 1921, Tondui 627 (US); Chiapas, Dept. Santa Rosa, Dec. 1X92, Heyde & Lux 4870 (G, NY, I 'S); Santa Rosa, May 1S92, Heyde & Lux 8019 in part (G). HoNuntAs: vicinity of Siguatepeque, Dept. Comayagua, Siandley 56082 (F, US). San Salvadoh: vicinity of Ateos, Dept. la Libertad, 17 April 1922, Standley 28826 (( !, US); Volcan de San Vicente, Dept. San Vicente, 7-8 March 1922, Standley 21486 (G, NY, US); Cerro de San Jacinto, near San Salvador, 8 Feb. 1922, Standley 20617 (O, US); San Jacinto, 5 April 1905, I elaseo 8848 (G, US); Volcan de San Salvador, 7 April 1922, Standley 22842 (US); vicinity of San Salvador, 1905, Renson 291 (NY, US); San Salvador, 1922, Calderon 729 (G, NY, US), 925 (G, US). Nicaragua: Casa Colorada and vicinity, south of Managua, 27 June 1923, Maxon, Harvey & Valentine 785(1 (NY, Vi>); Las Nubes and vicinity, south of Managua, 28 June 1923, Mason, Harvey & V (dentine 7478 (US). Costa Rica: without locality, 19 June 1S74, Kuntze 2109 (NY, US); without locality, April 1910, Worlhen (MPG); Los Ayotes, near Tilaran, Prov. Guanacaste, 21 Jan. 1926, Standley <«; Valerio 45 460 (US); Aguacate, Nov. 1846, Oersted 11822 (US); vicinity of San Jose, Feb. 1924, Standley 88298 (US); vicinity of La Palma, on the road to La Hondura, 17-18 July 1923, Maxon A Harvey 7951 (US); Cerro de Piedra Blanca, above Escasu, 31 Jan. 1924, Standley 82652 (US); near Cartago, 1845- X, Oerstnl 11824 (US); Cartago, Oct. 1SS7, Cooper 5890 (F, G, MBG, NY, US). Panama: fields along Rio Caldera above El Boquete, 5 Feb. 1918, Killip8510 (US). This is a widely distributed species with very distinctive habit and somewhat variable inflorescence. In some specimens the 1933] PERRY — NORTH AMERICAN SPECIES OF VERBENA 259 spikes appear to remain compact, in others they tend to elongate. Schauer used this difference to separate the forms pycnostachya and leptostachya, although he frankly admits the difficulty of distinguishing the two owing to the intermediate phases. V. affinis is characterized by a somewhat coarser floral pubescence; this, however, seems to be a variable feature and, as such, does not appear to merit more than passing mention. Ser. II. leptostachyae Schauer. Spikes slender and open or compact at anthesis, greatly elongating in fruit, solitary or in simple cymes or panicled. — North America. Key to the Species A. Spikes panicled at the apices of stems and branches, subtended chiefly by inconspicuous bracts; floral bracts not prominent. B. Spikes very slender, elongated and graceful, usually with remote fruits. C. Leaves 1-2-pinnatifid or 3-5-cleft or deeply incised. D. Corolla-limb not less than 3 mm. broad; nutlets 1.5 mm. or more long. E. Schizocarp a little longer than broad. F. Inflorescence usually densely glandular and somewhat viscid- pubescent 6. V. officinalis F. Inflorescence sparsely, if at all, glandular and strigillose 7. V. menthaefolia E. Schizocarp about twice as long as broad. G. Leaf-blades diverse in outline, the basal incised-dentate, the middle stem-leaves 1-2-pinnatifid, the upper sparingly toothed or entire; bracts one-half as long as the calyces at anthesis 8. V. Halei G. Leaf -blades similar in outline; bracts about as long as the calyces at anthesis 9. V. riparia D. Corolla-limb scarcely more than 1 mm. broad; nutlets 1.5 mm. or less long 10. V. Ehrenbergiana C. Leaves serrate or shallowly incised. H. Leaves subsessile or short-petioled with attenuate base. I. Stems hirsute-hispid; leaves mostly crenate-serrate; calyx hispidulous 11. V. Carolina I. Stems glabrous; leaves acutely serrate; calyx practically glabrous IS. V. longifolia H. Leaves obviously petioled with rounded-cuneate base. J. Fruiting-calyx spreading; calyx-lobes connivent; stigmatic surface subtended by two sterile style-lobes 1I+.Y . scabra J. Fruiting-calyx ascending; calyx-lobes not connivent; stigmatic surface subtended by one sterile style-lobe 15. V. urticifolia B. Spikes thicker or densely flowered, usually with contiguous fruits. 260 ANNALS OF THE MISSOURI HOTANICAL GARDEN [Vol. 20 K. Spikes subsessile or very short-peduncled, forming a congested in- florescence, not strict; corolla-limb 2 mm. broad 12. V. recta K. Spikes peduncled, usually not congested, strict; corolla-limb 3-4.5 mm. broad 16. V. hastata A. Spikes solitary or in 3's at the apices of stem and branches, or panicled and subtended by leafy bracts at the base; floral bracts prominent or not. L. Schizocarp readily separating into 4 nutlets at maturity. M. Leaves serrate-dentate or shallowly incised, or predominantly entire. N. Leaves serrate or serrate-dentate or shallowly incised; spikes with more or less contiguous fruits. (). Plants coarse, more or less densely hirsute-pubescent or hirsute- villous; leaves elongate-elliptical to ovate-orbicular; spikes stout at anthesis, 7-10 mm. broad. P. Leaves elongate-elliptical to ovate-acuminate, short-petiolate; corolla-limb 6-6 mm. broad. (J. Plants hirsute-hispidulous; spikes flexuous, elongated and open in fruit; bracts not exceeding the fruiting calyx; nutlets 2 mm. long, shallowly scrobiculate above, sulcate toward the base SI. V. macrodonta Q. Plants hirsute-pubescent; spike strict, dense in fruit; bracts surpassing the fruiting calyx; nutlets 2.5 mm. long, retic- ulate-scrobiculate above, striate toward the base 20. V. Mac Doug alii P. Leaves ovate-orbicular, sessile; corolla-limb 8-9 mm. broad. . 19. V. stricta (). Plants more slender, hirtellous or sparsely strigillose with short trichomes, canescent or not; leaves linear to narrowly elliptical or spathulate; spikes at anthesis 5 6 mm. broad. R. Leaves strigillose; inflorescence not glandular-pubescent. . . . R. Leaves hirtellous; inflorescence glandular-pubescent. 17. V. simplex S. Leaves hirtellous; rhachis more or less angulate; bracts lanceo- late-acuminate; corolla-limb 3-4 mm. broad. . .18. V. Orcuttiana S. Leaves hirtellous and canescent; rhachis scarcely angulate; bracts ovate-acuminate; corolla-limb 6-8 mm. broad. . . . 26b. V. neomexicana var. hirtella N. Leaves predominantly entire (mostly linear, the lower ones with a few salient teeth); spikes with more or less remote fruits. .27. V. perennis M. Leaves deeply incised-dentate or pinnatifid or 3-cleft. T. Spikes not essentially bracteose; flowers readily seen; corolla-limb 4-10 mm. broad. U. Leaf-blades, at least the lower ones, oblong-ovate or obtusely elliptic-ovate, not narrowly elongated, usually 3-cleft with the segments incised-dentate. V. Leaves petioled; pubescence various, but not coarsely hirsute; spikes usually stout, if slender not greatly elongated. W. Leaves not plicate, venation not noticeably whitish near the margin; spikes stout at anthesis, elongating or not in fruit. 1933] PERRY — NORTH AMERICAN SPECIES OF VERBENA 261 X. Plants pilose to hirsute-villous; leaves not scabrous above; spikes elongating in fruit; bracts not exceeding the calyx 22. V. prostrate X. Plants sparsely hirsute; leaves scabrous above; spikes occasionally elongating in fruit; bracts slightly sur- passing the calyx 23. V. robitsta W. Leaves more or less strongly plicate; venation noticeably whitish near the margin; spikes not stout at an thesis. . . . 25. V. plicata V. Leaves subsessile or at most very short-petioled; pubescence coarsely hirsute; spikes slender and greatly elongated. .24- V. xutha U. Leaf-blades, at least the lower ones, oblong-lanceolate to spathu- late, narrowly elongated, usually incised-pinnatifid or incised- dentate (excl. V. plicata). Y. Plants coarse with a low somewhat compact habit, canescent- hirsute; leaves subpinnatifid, contracted into a broadly margined semiamplexicaul or petiolar base. Z. Inflorescence somewhat glandular-hirsute; bracts lanceolate. 29. V. canescens Z. Inflorescence sparsely, if at all, glandular and densely hirsute; bracts ovate, abruptly acuminate 29a. V. canescens var. Roemeriana Y. Plants more slender with a taller and open habit, hirsute to canescent-hirtellous; leaves .pinnately cleft to incised- dentate, with a narrowly margined petiolar base. AA. Corolla-limb 4 mm. broad; commissural faces extending to the tip of the nutlet 26. V. neomexicana AA. Corolla-limb 6-10 mm. broad; commissural faces scarcely extending to the tip of the nutlet. BB. Plant more or less hirsute; leaves pinnately cleft; bracts lanceolate. . 26a. V. neomexicana var. xylopoda BB. Plants canescent-hirtellous; leaves dentate or somewhat more deeply incised; bracts ovate 26b. V. neomexicana var. hirtella T. Spikes usually bracteose with somewhat foliaceous bracts at the base of the spike; flowers inconspicuous; corolla-limb 2.5-3 mm. broad (larger in V. plicata). CC. Plants delicate; leaves incised-pinnatifid to pinnately cleft, not pli- cate; inflorescence glandular-hirtellous; nutlets 1.5-2 mm. long, scrobiculate practically to the base 28. V. gracilis CC. Plants coarse; leaves incised-dentate to subpinnatifid or 3-cleft, pli- cate or not; inflorescence hirsute or pubescent-hirsute; nutlets 2-2.5 mm. long, not scrobiculate to the base. DD. Leaves more or less strongly plicate, conspicuously whitish- veined near the margin 25. V. plicata DD. Leaves not plicate nor conspicuously whitish-veined near the margin. EE. Plants hirsute; leaves spreading-hirsute; nutlets raised- reticulate above, striate below. FF. Leaves with a subpetiolar or semiamplexicaul base; bracts 262 ANNALS OF THE MISSOURI BOTANICAL GARDEN | Vol. 20 ovate (a little longer than the flowers), abruptly acumin- ate, ascending 29a. V. canescens var. Roerneriana FF. Leaves narrowed into a margined petiole; bracts linear- lanceolate (much longer than the flowers), usually re- flexed in age 31. V. bracteata KE. Plants hirsute-pubescent; leaves appressed-hirsute; nutlets only faintly reticulate or essentially smooth 30. V. subidigera L. Sohisooarp tardily separating into 4 nutlets at maturity 32. V. carnea 6. V. officinalis L. Sp. PI. 20. 1753. V. spuria L. I.e. V. darning ensis Urb. Symb. Ant. 5: 484. 1908. Stems ascending or erect, branched, glabrous or nearly so; leaves 2-7 cm. long, strigillose on both surfaces; basal and lower stem-leaves more or less ovate, narrowed below into a petiole, 1-2-pin natifid or 3-5-cleft with parts incised; upper leaves similar but smaller and less divided; spikes paniculately disposed or in 3's or solitary, slender and elongate; bracts usually about half as long as the calyx; calyx 2-2.5 mm. long, glandular-pubescent, subtruncate, teeth minute; corolla-tube a little longer than the calyx; corolla-limb about 4 mm, broad, segments more or less rounded; nutlets trigonous, barely 2 mm. long, strongly striate, slightly reticulate above; commissural faces muricate. Distribution: Europe; introduced in waste places in eastern North America and West Indies. Specimen* examined: Massachusetts: Rowley, Oakes (G). Rhode Island: Warwick Neck, 1848, Thurber (G). New Yowl: Brooklyn, 13 Sept. 1879, von Schrenk (MBG). New Jersey: ballast, Camden, 30 Aug. 1874, Parker (G); roadside, Longacoming, Camden Co., 23 July 1867, Parker (G); Cold Spring, Cape May Co., 30 Aug. 1917, Gershoy 583 (G). Pennhvlvania: Philadelphia, 1844, Lea (MBG); Lancaster, 21 Aug. 1861, Porter (G); banks of Susquehanna, Harrisburg, Oct. 1852, Porter (G); York Furnace, York Co., 28 June 1899, MacElwee 873 (MBG). Delaware: Wilmington, 1845, Tatnall (G). District of Columbia: vicinity of Washington, 21} June 1878, Ward (MBG). Virginia: Parksley, 11 Sept. 1902, Norton (MBG); Bedford Co., 30 June 1870, Curtiss (MBG); Marion, 1892, Small (MB< J). North Carolina: without data, Curtis (MBG); waste grounds, Moyock, 1 July 1922, Randolph 587 (G); dry sandy bank about, 3 km. west of Plymouth, 4 July 1922, Randolph 645 (G); Clarkton, 23 June 1897, Bilimore Herbarium 4762 (G, NY); Cranberry Forge, July 1895, F. Wislizenus 1214 (MBG). South Carolina: streets in Gnmiteville, 23 May 1899, Eggeri (MBG). Georoia: Rome, Chapman (MBG). 1933| PERRY — NORTH AMERICAN SPECIES OF VERBENA 263 Florida: without data, Rugel 121 (F, MBG). Alabama: Collinsville, 29 July 1897, Eggert (MBG); Attalla, 9 July 1898, Eggert (MBG). Louisiana: Port Eads, 22 Aug. 1900, Tracy & Lloyd (G, MBG, NY). Tennessee: along the banks of Doe River, Carter Co., 16-17 July 1891, Small & Heller 484 (G, MBG); Knoxville, July 1893, Ruth (MBG); Hollow Rock, Carroll Co., 5 Aug. 1897, Eggert (F, MBG). West Indies: Bermuda: Hamilton Parish, 11 July 1905, Moore 2989a (G); Flatts, 31 Aug.-20 Sept. 1905, Brown & Button 28 (ANSP, G, NY); Flatts, 3 Aug. 1915, Collins 267 (G, NY); Bailey's Bay, 10 Feb.-9 March 1908, Brown 492 (ANSP, G, NY); St. George's, 18 Jan. 1912, Robinson 113 (G). Santo Domingo: Angostura del Rio Yaqui, 8 May 1887, Eggers 1828 (NY), type collection of V. domingensis ; Culo de Maco, Prov. Azua, Aug. 1912, Fuertes 1856 (NY); Loma Rosilla, Prov. de la Vega, July 1912, Fuertes 1771 (NY); vicinity of Mission, Fonds Varettes, 12 April-4 May 1920, Leonard 3939 (G, NY). Cuba: without data, Wright 3658 (G); Vecindad de Vento, May 1906, Baker 2591 (NY, P); Playa de Marianao, 29 March 1911, Britton & Cowell 10326 (NY); near Playa de Marianao, Leon (NY), Leon & Edmunds (NY); cultivated field, Campo Florido, Prov. Havana, 13 March 1905, Curtiss 677 (ANSP, G, MBG, NY). On the whole, the specimens from Santo Domingo and Cuba differ from the typical V. officinalis in their slender and more elongate habit; the inflorescence is scarcely as glandular, the flowers are smaller, and the nutlets often do not exceed 1.5 mm. in length. Nevertheless, the Cuban specimens vary greatly in size, and Curtiss 677 is hardly separable from typical V. officinalis. Since many of the specimens are rather poor, it appears probable that they may very well represent an impoverished condition. Urban himself was somewhat uncertain of the status of his species as he appended the following note in a later publication: "An re vera a formis V. officinalis L. separanda?" 7. V. menthaefolia Benth. PI. Hartw. 21. 1839. V. setosa Mart. & Gal. Bull. Acad. Brux. II 2 : 321. 1844. Stems decumbent or ascending, branched, sparsely and minutely hispidulous; leaves ovate, tapering at base into a margined petiole, 3-6 cm. long, deeply cleft or subincised with divisions remotely serrate-dentate, strigillose on both surfaces, somewhat pustulate above; spikes panicled, slender, elongate, compact only at the apex; bracts ovate-lanceolate, acuminate, variable in length, usually shorter than the calyx, ciliate and sparsely strigillose; calyx 2.5-3 mm. long, strigillose, sparsely (if at all) glandular, teeth minute; corolla-tube only slightly longer (Vol. 20 2G4 ANNALS OF THE MISSOURI BOTANICAL (JARDEN than the calyx; corolla-limb about 6 mm. broad, segments more or less truncate; nutlets trigonous with convex back, 2-2.5 mm. long, striate, raised-reticulate above; commissural faces muricate. Distribution: Arizona to southern California, northwest Mexico to Oaxaca. Specimens examined: Arizona : north of Yuma, 26 April 1906, Jones (P). California: San Diego, May 1852, Thurber 555 (G), June 1875, Palmer 808 (MBG), 5 May 1903, Abrams 8406 (G, MBG, NY, P), 6 July 1915, Macbride & Payson 781 (G), 17 April 1 1 1 1 8 , Carlson (G), in canyons, Aug. 1918, Spencer 971 (G, P), waysides, 31 Oct. 1919, Spencer 11,14 (G, P); Otay Creek, San Diego Co., 27 March 1923, Peiraon 8379 (P). Mexico: Loweii California: Salton River Crossing, 27 April 1894, Schoenfeldt 2915 (G, NY, US); sandy roadside, Tia Juana, 1 Feb. 1920, linrtram (ANSP); ranch, 46 km. southwest of Tia Juana, 13 April 1925, Jones (!'); base of Cucopa Mountains, 6 April 1905, MacDougal 158 (NY). Sonoha: vicinity of Alamos, 16 March 1910, Rose, Standley & Russell 12984 (US); vicinity of Hermosillo, 6 March 1910, Rose, Standley & Russell 12451 (US); vicinity of Navojoa, 21 March 1910, Rose, Standley cfc Russell 18130 (US). Sinaloa: collection of 1921, Ortega 4215 (US); vicinity of Fuertc, 25 March 1910, Rose, Standby A Russell 13447 (MS); vicinity of San Bias, 24 March 1910, Rose, Standley A Russell 13422 (NY, US); Topolobampo, 15-25 Sept. 1897, Palmer 268 (US); near Plomosas, 18 July 1897, Rose 1763 (US). Chihuahua: valley of the San Pedro, Ortiz, 11 April 1887, Pringle 1599 (MBG). Coahuila: near Saltillo, 7 May 1848, Gregg 11 (MBG); near Saltillo, 14 July 1848, Gregg 265 (( 1, MBG); vicinity of Saltillo, May 18«i8, Pidrner 191 (G, MBG, NY, US); vicinity of Buena Vista, 24 July 1848, Gregg 276 (MBG); valley of Parras, 11 April 1847, Gregg 406 (MBG, NY); San Lorenzo de Laguna, about 120 km. southwest of Parras, 1-10 May 1880, 1'ulmer 1042 (ANSP, G, US). DuRANCio: near El Salto, 12 July 1898, Nelson 4577 (MBG, US); bottom-lands and ravines, Durango, April-Nov. 1896, Palmer 153 (F, G, MBG, NY, US), 856 (G, MBG, NY, US). San Liiih PoTOfli: Alvarez, 28 Feb.-3 Oct. 1902, Palmer 141 in part (F, G, MBG, NY, US); San Luis Potosi, 1878, Parry & Palmer 717 (G). Aguahualientes: near Aguas Calientes, 10 Oct. 1903, Rose & Painter 7799 (US). Guanajuato: margin of stream, Sirena Mountain, 1894, Duges (G); Leon, 1839, Hartweg 175 (K type, NY). Quehetaho: vicinity of Queretaro, 1912, Basile 99 (US); Queretaro, 1910-13, Arsbne t fc Agniel 10242 (F, G, MBG, US); Queretaro, July 1914, Arsene 9998 (US). Vera Cruz: Maltrata, 20 Aug. 1891, Seaton 7 (V, G, US); Huatusco, 1857, Mohr (US). HinALQo: Sierra de Pachuca, 20-24 July 1905, Rose, Painter & Rose 8753 (G, NY, US); Nopala, Aug. 1913, Salazar (US); Moran, 1840, Galeotti 778 (K), tyi-e collec- tion of V. setosa. Mexico: near Tlalnepantla, 6 July 1905, Rose, Painter A Rose 8382 (G, US); Lomas de Santa Fe, July 1928, Lyonnet (US); Valley of Mexico, 1865-66, Iiourgeau 860 (US), 547 (G); Valley of Mexico, 27 June 1901, Pringle 8584 (ANSP, F, G, MBG, 1933] PERRY — NORTH AMERICAN SPECIES OF VERBENA 265 NY, P, US); mountains between Toluca and Mexico City, 28 June 1910, Rusby 181 (NY). Morelob: La Cascada, 29 May 1901, Pringle 9529 in part (F, US). Michoacan: Morelia-Rincon, Arsene 2798 (G); east of Maravatio, 30 April 1844, Gregg 823 (MBG). Oaxaca: mountains, San Juan del Estado, 18 June 1894, L. C. Smith 27 (G); Sierra de San Felipe, 20 Aug. 1894, Pringle 5715 (G); between Coixtlahuaca and Tamazulapam, 12 Nov. 1894, Nelson 1943 (US); between Las Sedas and Salome, 30 Aug. 1921, Conzatti 4207 (US). In the specimens cited from California south including Sinaloa, the inflorescence is more densely strigillose than in the collections from the southern part of Mexico, the calyces are about 1 mm. longer, with teeth strongly unequal and the subtending bracts often as long as the calyces. Although this apparently indigenous species has been known generally as V. officinalis, it has somewhat harsher pubescence and is scarcely, if at all, glandular. The fruiting calyx tends to be connivent, concealing the apex of the schizocarp rather than open and disclosing it. Perhaps these are differences only of degree and may be merely variations of V. officinalis; nevertheless, for the present it seems preferable to retain the name V. menthaefolia for the American representative. 8. V. Halei Small, Bull. Torr. Bot. Club 25: 617. 1898. V. leucanthemifolia Greene, Pittonia 5: 135. 1903. Stems usually several from a woody base, erect, ascendingly branched, glabrous or strigillose above; leaves 3-10 cm. long, strigillose on both surfaces, diverse in outline — the basal and lower stem-leaves oblong to ovate, tapering into a petiole ap- proximately as long as the blade, irregularly dentate or incised; the middle stem-leaves 1-2-pinnatifid with shorter petioles; the upper sparingly dentate or entire; spikes paniculately disposed, slender and elongate; bracts about one-half as long as the calyx, appressed, ciliate; fruiting calyx 3-3.5 mm. long, strigillose, subtruncate, the broad nerves terminating in unequal subulate teeth; corolla-tube scarcely longer than the calyx; corolla-limb 6-7 mm. broad, segments retuse; nutlets trigonous, approximately 2.5 mm. long, usually prominently striate, raised-reticulate at apex; commissural faces muricate. Distribution: Alabama to Texas, Mexico. Alabama: without data, Buckley (MBG); Frascati, 11 May 1904, Deweys (G). 260 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 20 Mississippi: cultivated grounds near coast, May 1859, Hilgard (MBG); Ocean Springs, 14 Sept. 1891, Seymour 50 (G); Ocean Springs, June 1892, Skehan 109 (G, MBG), 46 (, Chandler (G, MBG); Sierra Blanca, 11 April 1930, Jones 962*9 in part (MBG). Mexico: Tamaulipas: Matamoros, Berlandier 1511, 8016 (G, MBG); vicinity of Tampico, 1-31 Jan. 1910, Palmer 78 (F, G, MBG, NY, US). V species closely related to V. menthaefolia and V. officinalis, 1933] PERRY — NORTH AMERICAN SPECIES OF VERBENA 267 but readily distinguished by the diverse outline of the and the somewhat more slender achenes. 9. V. riparia Raf. ex Small & Heller, Mem. Torr. Bot. Club 3:12. 1892. ? V. hastata var. p oblongifolia Nutt. Gen. 2: 40. 1818. V. urticifolia var. riparia Britton, Mem. Torr. Bot. Club 5: 276. 1894. Stems 6-15 dm. tall, erect, sparsely pubescent or glabrate, widely branched; leaves oblong to ovate, 4-12 cm. long, petioled, pinnatifid or nearly tripartite toward the base, sparsely strigillose on both surfaces, venation prominent beneath ; spikes paniculately disposed, slender, elongate; bracts lanceolate-ovate, as long as the calyx at an thesis, acuminate ; calyx 3 mm. long, minutely glandular- pubescent, subtruncate, teeth minute; corolla-tube only slightly longer than the calyx, puberulent without; corolla-limb 3.5 mm. broad; segments more or less rounded, the middle posterior one emarginate; nutlets oblong, 2-2.5 mm. long. Distribution: New Jersey and Virginia to North Carolina (ace. to Small). Specimens examined: Virginia: Marion, Smyth Co., 1 July 1892, Small (P); Marion, Smyth Co., 6 July 1892, Small (G, MBG). North Carolina: near Globe, Caldwell Co., 3 July 1891, Small & Heller (F); near falls of Yadkin, Stanley Co., 18 Aug. 1891, Small & Heller (F). The material at hand is too scanty and too immature to give many clues to the probable relationship of this species. It would seem as nearly related to V. officinalis as to either V. hastata or V. urticifolia. It is characterized by pinnatifid or tripartite leaves, minutely glandular-pubescent inflorescence, and fruit about twice as long as thick. 10. V. Ehrenbergiana Schauer in DC. Prodr. 11 : 548. 1847. Stem erect, branched, hirsute; leaves trifid with lateral lobes small, somewhat ovate with cuneate base narrowed into a mar- gined petiole, attenuate at apex, 4-8 (-10) cm. long, coarsely serrate-dentate, strigillose above, hirtellous below, also somewhat paler and prominently veined; spikes paniculately disposed, very slender, remotely flowered; bracts ovate, about one-half as long as the calyx or shorter, subulate, ciliate; fruiting calyx [VOL - 20 268 ANNALS OF THE MISSOURI HOTANICAL GARDEN about 1 .5 mm. long, strigillose, lobes very short, obtuse, mucro- nate; corolla inconspicuous, limb about 1 mm. broad; nutlets trigonous with convex back, 1 mm. long; commissural faces Pringl meeting sharply at right angles, almost smooth Distribution: Mexico. Specimens examined: Mexico: COAHTJILA: 8altillo, July 1880, Palmer 2037 (O). Nuevo Leon: moist places near Monterey, July 1888, MUG, NY, IIS). San Luis Potosi: Tancanhuitz, Feb. 1888, Sdtr 722 (G, US); Rio Verde, 17 Nov. 1910, Ormtt 6 ',23 (MBG); Bagre, Minas de San Rafael, July 1911, Purvus 5L51 (F, G, MBG, NY, US). Hidalgo: Cazadero, April 1841, Licbmann 1 1335 (US). Vera Cituz: Wartenborg, near Tantoyuca, lN. r >8, E rvendberg 153 (ANSP, G). Pttxbla: Pahuatlan, 12 July 19K5, Salazar (US). Mexico: near Los Reyes, Ehrcnberg 713 (Bot. Mus. Berl.-Dahl. type, MBG phot.). In general habit this species somewhat approaches V. Carolina. It differs, however, in its tripartite leaves, smaller flowers, and fruits. The collection from Saltillo is much more densely hirsute on all parts, but aDDarentlv is cnnsnpmhn 11. V. Carolina L. Syst. ed. 10, 852. 1759; Sp. PI. ed. 2, 29. 1762. V. caroliniana Willd. Sp. PL 1 : 119. 1798. V. polystachya IIBK. Nov. Gen. et Sp. 2: 274. 1818. V. biscmila HBK. I. c. 275. V. veronicaefolia IIBK. I. c. V. hirsula Mart. & Gal. in Bull. Acad. Brux. II 2 : 321. 1844. V. mollu Mart. & Gal. I. c. 323. V. paucifolia Mart. & Gal. /. c. 324, as V. paucijlora M. & G. in Walp. Hep. 6: 087. 1846-47. V. caroliniana forma or var. polystachya (Kunth) Loes. in Fedde, Rep. Sp. Nov. Veg. 9: 3G2. 1911. Stems erect or ascending, usually solitary, branched, hirsute- hispid; leaves lanceolate-oblong or somewhat elliptical, 3-8(-12) cm. long, obtusish or acute, subsessile or narrowed into a short petiole, coarsely crenate-serrate, strigose and somewhat pustulate above, hirsute-strigose especially along the midrib and veins beneath; spikes paniculately disposed, slender, open in fruit; bracts ovate, varying in length, usuallv about one-half as Inner n* 1933] PERRY — NORTH AMERICAN SPECIES OF VERBENA 269 the calyx or shorter, acuminate-subulate, somewhat spreading, ciliate; fruiting calyx approximately 2 mm. long, with the obtuse mucronate lobes connivent, hispidulous; corolla-tube scarcely exserted; corolla-limb inconspicuous, about 2 mm. broad; mature schizocarp inclosed by calyx; nutlets trigonous with convex back, hardly 1.5 mm. long, faintly striate; commissural faces meeting at right angles, almost smooth. Distribution: Arizona and Mexico to Salvador, Central America. Specimens examined: Arizona: Wilgus Ranch, Chiricahua Mountains, 12 Oct. 1907, Blumer 1783 (G, MBG, NY, US); Cave Creek, Chiricahua Mountains, 2 Sept. 1929, Harrison & Kearney 6144 (US) ; Santa Rita Mountains, 15 July 1881, Pringle (G); Fort Huachuca, July 1893, Wilcox (NY); Huachuca Mountains, 4 Sept. 1903, Jones (P, US); Ramsay Canyon, Huachuca Mountains, 30 Sept. 1929, Jones 25006 (G, NY). Nevada: Carson Valley, Aug. 1872, Lemmon 3075 (G). Mexico: without data, Graham (G); collection of 1844, Galeotti 795 A (G, US). Lower California: Sierra de San Francisquito, 1 Oct. 1899, Brandegee (NY). Sonora: Santa Cruz, 22 Oct. 1893, Mearns 2627 (US); Cochuto, 1 Oct. 1890, Hartman 94 (G, NY, US); La Cruz de los Canadas, 3 Nov. 1890, Lloyd 448 (G). Sinaloa: near Colomas, foothills of Sierra Madre, 15 July 1897, Rose 1677 (US); Santa Lucia, 1919, Dehesa 1551 (US). Chihuahua: Norogachi, Aug.-Nov. 1885, Palmer 364 (ANSP, G, US); San Diego Canyon, Sierra Madre Mountains, 16 Sept. 1903, Jones (P); Guayanopa Canyon, Sierra Madre Mountains, 23 Sept. 1903, Jones (P). Durango: rich bottom-lands, vicinity of Durango, Apr-No v. 1896, Palmer 339 (F, G, MBG, NY, US). Nayarit: west of Tepic, 31 May 1849, Gregg 1001 (MBG); Tepic, 5 Jan.-6 Feb. 1892, Palmer 2057 (US). Jalisco: Cuidad Guzman, 14 Oct. 1921, Kempton & Collins (US). Guanajuato: Guanajuato, 1880, Duges (G). Hidalgo: Real del Monte, 15 Sept. 1910, Clokey 1865 (MBG); near Moran, Humboldt & Bonpland (Par. type of V. veronicaefolia, MBG phot.). Vera Cruz: Sierra Templada in general, June-Oct. 1840, Galeotti 735 (K), type collection of V. hirsuta; Orizaba, 1853, Mueller (NY); Orizaba, 1855, Mueller 766, 887 (NY); Orizaba, 1855-7, Botteri 180 (F, G, MBG, US); Orizaba, 1857, Mohr (US); Jalapa, Schiede 88 (NY); Jalapa, Jan. 1894, C. L. Smith 1737 (G). Puebla: region of Atoyac, near Puebla, 15 July 1909, Nicholas (US); Tezuitlan, 9 June 1910, Orcutt 4045 (F). Mexico: near city of Mexico, Humboldt & Bonpland (Far. type of V. biserrata 9 MBG phot.); valley of Mexico to Tizapan, 19 Aug. 1865-66, Bourgeau 119 (G, US); Chalco region, 4 Oct. 1921, Kempton & Collins (US); near Tlalpam, 1905, Rose, Painter & Rose 8496 (US); Tlalpam, 9 Aug. 1910, Orcutt 3488 (F, US); Eslava, 15 June 1901, Pringle 9312 (F, G, MBG, NY, US); near Toluca, 18 Sept. 1889, Pringle 2813 (F, G). Morelos: fields, La Cascada, 29 May 1901, Pringle 9529 in part (G, MBG, NY); Toro, 5 Aug. 1924, Fisher (F, MBG); Tepoztlan, autumn 1926, R. Redfield 4 (US). Michoacan: Morelia, Coronilla, 8 Aug. 1909, Arsbne 3000 (US); Quinceo, near [Vol. 20 270 ANNALS OF THE MISSOURI BOTANICAL GARDEN Morelia, 1 1 Nov. 1009, Arscne (US); on slope of volcanic mountain, Xorullo, Hum- boldt (.{'. Bonpland (Par. tyi>k of V. poiystachya, MBG phot.). Colima: Colima, 9 Jan.-6 Feb. 1891, Palmer 1156 (G, NY, US). Oaxaca: mountains near Oaxaca, June-Oct. IS40, Caleotti 737 (K), type collec- tion of V. mollis; Sierra de San Felipe, 12 Sept. 1804, Pringle 489$ (ANSP, G, MBG, NY, US); Sierra de San Felipe, 2 Oct. 1S04, C. L. Smith 224 (MBG, NY, US); vicinity of La Parada, 10 Aug. 1S04, Nelson 102 I (US); Cuyamecalco, 5 Sept. 1805, L. C. Smith 634 (G). Centkai, America: Guatemala: uncultivated places, Oct. 1865, Bernoulli 127 (NY); without locality, Sept. 1027, Morales 786 (US); without locality, 1892, Heyde 120, 530, 477 (US); S Siguan, Dept. Quiche, April 1802, Heyde 5944 (F, US). Salvadok: vicinity <.f A paste pe< pie, Dept. San Vincente, 4 March 1022, Standley 21331 ((!, MUG, US); vicinity of San Salvador, h'enson 175 (NY, US); Plazuela de Aculhaca, San Salvador, Dec. 1006, Velasco 8999 (US); Amatepeque Hill, near San Salvador, 2 Pel.. 1007, Vittier 1909 (US); San Salvador, 1022, Calderon 794 (G, V^); vicinity of San Salvador, 30 March-24 April 1022, Standley 22414. (G, NY, US); vicinity of Santa Tecla, Dept La Libertad, 10 April 1922, Standley 23083 (G, US); Finca San Nicolas, 1923, Choussy 36 (US). This species is probably most nearly related to V. urticifolia and V. 8cabra, but is easily set apart from them by its subsessile or very short-pet iolate leaves; moreover, their areas of distribu- tion are practically distinct. The collections cited appear to be conspecific, notwithstanding the fact that they vary greatly in size, serration of leaves, and length of floral bracts. The speci- mens from the northern part of the range are slightly coarser, with leaves more remotely crenate-serrate, yet they match very 4892, from the Sieri the hitter, however, In pubescent on the under surface. The specimens from Guatemala and Sal- vador have somewhat crowded spikes and are, as a whole, smaller in every way and more densely hispidulous. However, since most of the collections were made from December to Febru- compact habit might be graph 1933] PERRY — NORTH AMERICAN SPECIES OF VERBENA 271 A specimen labelled "Inter Tallahassee et St, Marks, Florida, legit Rugel, April- June 1843 " appears in the herbarium of the Missouri Botanical Garden. As this is the only representative of V. Carolina seen from Florida, and as it is so far from the natural range of the species, it would appear to be an error in labelling. Another collection worthy of mention is Nelson 752. It is almost glabrous, and the lobes of the corolla are emarginate, a rather unusual feature in this series. The lack of pubescence suggests V. longifolia, but unfortunately none of the available material of the species is in sufficiently good condition to reveal the character of the corolla. Although the general habit of the collection is that of V. Carolina, it is rather doubtfully con- specific, 12. V. recta HBK. Nov. Gen. et Sp. 2: 277. 1818. V. caroliniana forma or var. recta (Kunth) Loes. in Fedde, Rep. Sp. Nov. Veg. 9:362. 1911. Stem erect, branched, hispidulous, reddish; leaves lanceolate- oblong to ovate with cuneate base narrowed into a short margined petiole, 4-8 cm. long, acute, serrate-dentate, reticulately veined, strigillose on both surfaces, minutely pustulate above; spikes numerous, congested at the apices of stems and branches, short- cylindrical, usually dense but not strict; bracts a little shorter than the calyx, ovate-lanceolate, acuminate, hispidulous; fruiting calyx about 2 mm. long, hispidulous, lobes short, obtuse, mu- cronate, commonly connivent; corolla-tube scarcely exserted; corolla-limb inconspicuous; nutlets trigonous with convex back, 1.5-2 mm. long, faintly striate; commissural faces meeting at right angles, practically smooth. Distribution: Mexico. Specimens examined: Mexico: Hidalgo: Real del Monte, 12 July 1913, Salazar (US); between Pachuca and Cerro Ventoso, Humboldt & Bonpland Jfi66 (par. type, MBG phot.). Puebla: Huitzuilzilapam, 9 June 1910, Orcutt 3950 (F). Mexico: Cima Station, 30 Aug. 1905, Pringle 13597 (G, US); Toluca, Berlandier 1222 (US). Morelos: Tres Marias, 4 July 1901, Rose & Hay 5310 (US); Toro, 5 Aug. 1924, Fisher 820 (MBG, US). Oaxaca: vicinity of Cerro San Felipe, 1894, Nelson 1105 (US); Sierra de San Felipe, 9 Aug. 1894, Pringle 4769 (ANSP, F, G, MBG, NY, US). 272 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 20 Verbena recta, generally determined in the material at hand as "aff. V. polystachya IIBK.," is easily distinguished from this closely related species by the short hispidulous pubescence, the commonly broader leaves, and the compact cylindrical congested spikes. The floral characters of both are similar, although generally the fruiting calyx of V. Carolina is more acutely ovoid and the subtending bract is much sharper at the apex. 13. V. longifolia Mart. & Gal. Bull. Acad. Brux. II 2 : 323. 1844. Stems erect, tall, glabrous, obtusely 4-angled; leaves lanceolate to elongate-elliptic, 10 12 cm. long, the upper somewhat smaller, decussate, short-pet iolate, acutely serrate from below the middle to the apex, appressed-pubescent or very short-strigillose on both surfaces, venation prominent beneath; spikes paniculately disposed, elongate, slender, open in fruit, glabrous; bracts ovate, about one-half as long as the calyx, acute-acuminate, minutely cilia te ; calyx 2 mm. long, with the obtuse lobes connivent over the schizocarp, practically glabrous; corolla-tube scarcely pro- truding beyond the calyx; corolla-limb inconspicuous; nutlets trigonous, hardly 1.5 mm. long, smooth or faintly striate; com- missural faces smooth. Distribution: southern Mexico. Specimens examined: Mexico: Veua Ciuiz: Colipa, March 1841, Lidmann 11818 (US); near Chila, April 1888, Seler 724 (G, US). Pub B LA: in vicinity of San Luis Tultitlanapa, July 1908, Purpus SJfiG (F, G, MBG, NY, 1 S). MORE LOB: San Anion, near Cuernavaea, 11 Oct. 1904, Seler 4-194 (G, US); Cuer- navaea, 28 ( >ct. 1905, Seler J,84? (G, US). Oaxaca: Tehuantepec, 10 April 1910, OrcuttSS2l (F, MBG, US). A rather singular species combining the foliar characters of V. litoraUs with the inflorescence characters of V. Carolina. in. 2: 2. 1798, not V. scabra pi 5/ h fig.2. 1840-41. more cm hispidulous; leaves ovate to elongate-ovate, 3 1 ()(— 13) 2.5-5 cm. broad, petiolate, serrate-dentate, acute or obtu 1933] PERRY — NORTH AMERICAN SPECIES OF VERBENA 273 scabrous and commonly strigillose above, less scabrous and some- what paler beneath, also hispidulous along the veins; spikes paniculately disposed, slender, pedunculate, copiously and closely flowered; bracts ovate-acuminate, about half as long as the calyx, hispidulous; fruiting calyx 2.5-3 mm. long, hispidulous, ovoid, with the somewhat unequal lobes acutely connivent, diverging from the rhachis of the spike 45° or more; corolla-tube scarcely, if at all, longer than the calyx; corolla-limb about 2 mm. wide, lobes obtuse; anthers glandless; stigmatic surface midway between two almost equal obtusish sterile lobes; nutlets trigonous, 1-1.3 mm. long, faintly striate, reticulate above; commissural faces extending to the top of the schizocarp, meeting sharply at right angles, muriculate. Distribution: North Carolina, Florida west to California, south into northwestern Mexico; West Indies. Specimens examined: North Carolina: moist rich soil, open swamp land, Wilmington, 25 July 1922, Randolph 1012 (G). Florida: without locality, 1842-49, Rugel 156 (F, MBG); common in everglades, 27 Aug. 1925, O'Neill (MBG); vicinity of Mayport and Jacksonville, 1870-76, Keeler (NY); near Jacksonville, 25 June-13 July 1894, Curtiss 5111 (MBG, NY); Lake City, 21 July 1893, Qaaintance (MBG); vicinity of Eustis, Lake Co., 1-15 July 1894, Nash 1248 (G, MBG, NY); low ground, Orange Co., 22 July 1902, Fredholm 54W (G, MBG, P, US); open low ground, near Lake Okeechobee, 20 May 1925, E. J. Palmer 27462 (MBG); east shore of Lake Okeechobee, 11-25 Nov. 1913, Small & Small 4337 (NY); east shore of Observation Island, 11-25 Nov. 1913, Small & Small 4403 (NY); hammocks along Taylor Creek, 11-25 Nov. 1913, Small & Small 4341 (NY); Fort Lauderdale, 19-25 Nov. 1903, Small & Carter 1072 (F, NY); moist sandy soil, Hollywood, Broward Co., 10 Feb. 1930, Moldenke 591, 599a (MBG, NY); everglades west of Miami, 3 Nov.-7 Dec. 1912, Small 4020 (NY); hammocks near Miami River, 26 Nov.-20 Dec. 1913, Small & Small 4520 (NY); Biscayne Bay-Indian River, Palmer 397 (G, MBG); Myers, Lee Co., July- Aug. 1900, Hitchcock 269 (MBG); sandy ditch, south of Estero, Lee Co., 14 April 1930, Moldenke 972 (MBG, NY); Sanibel Island, 18 May 1901, Tracy (NY); Manatee, 8 May 1900, Tracy 6652 (G, MBG, NY); Hernando Co., June-July 1898, Hitchcock (MBG); near Tallahassee, Berg (NY); shore of Dog Island Sound, near Lanark, 26 July 1920, Harper 242 (NY, US); streets of Apa[lachicola], Chapman (MBG). Mississippi: Scranton, Jackson Co., 5-6 Aug. 1896, Pollard 1191 (G, MBG, NY); Ocean Springs, 13 Aug. 1895, Skehan (MBG). Louisiana: South Pass, 20 Aug. 1900, Tracy & Lloyd 22 (G, MBG); New Orleans, Riddell (G); New Orleans, Drummond (NY); New Orleans, 21 Oct. 1885, Joor (MBG). Texas: without locality, July 1851, Lindheimer 618 (MBG); Beaumont, Jefferson Co., 11 Sept. 1916, E. J. Palmer 10692 (MBG); Beaumont, Sept. 1904, Kirn 2139 (P); Houston, July 1842, Lindheimer (MBG); Comanche Springs, New Braunfels, 274 ANNALS OF THE MISSOURI liOTANICAl, (JARDEN [Vol 20 etc., Aug. 1851, Limlhnmvr 1077 (G, MBG, NY); Leon River, Oct. 1850, Bigelow (G); San Marcos and vicinity, 1898, Stanfield (NY); New Braunfels, 1847, Lind- hcimer (MBG). Aki/ona: north of Rice, Apache Reservation, 4 Oct. 1927, Harrison 4897 (US); Santa Crui River, Tucson, 11 May 18X1, Pringle (G); banks of Santa Cruz River, near Tucson, 18 July 1884, Pringle (ANSP, F, ( !, NY, P, US). California: without locality, Wallace (G); Gienega, on Santa Monica R. R., Lyon 6 (G); Los Angeles, 1892, Davidson (G); near Pasadena, 17 Dec. 1892, Mc- Clatchie (NY); San Bernardino, Oct. 1891, Parish (MBG), 5888 (NY); San Bernar- dino Valley, Parish & Parish 101,8 i F, MBG); San Bernardino Valley, 25 July 1909, Parish 711,9 (P); 3 km. south of San Bernardino, 2 Sept. 1924, Johnston (P); Upland, July 1917, Johnston (P); 8 km. southeast of Chino, Mum & Johnston 11289 (P); CooomoiO Mountains, Aug. 1881, Parish & Parish 1111,8 (G). Mexico: LOWSR California: San Tomas, 15 July 1886, Orcutl 1802 (MBG). CoAHUlLA: Soledad, a section of low mountains with few oaks, 40 km. southwest from Monoclova, 9-19 Sept. 1880, Palmer 101,0 (ANSP, US); San Lorenzo de Laguna, 120 km. southwest of Parras, May 1XS0, I'almrr 101,0 (G). West Indies: Cuba: without locality, 1865, Wright 8669 (G); Santa Rosalia, 2 Aug. 1895, Combs 889 (( 1, MBG, NY); valley of San Juan, vicinity of Matnnzas, 14 March 1903, Britton, Britton & Shafer 292 (NY); Havana, 15 Oct. 1908, Leon (185 (NY); Havana, 21 April 1919, Leon & Edmunds 8719 (NY); Rio Arimao, 22 March 1910, Britton A Wilson 6771 (NY); Isle of Pines, 8 March 1916, Britton, Wilson & Lam 15256 (NY); Pinar del Rio, 7 April 1024, Krig 8187 (NY). Jamaica: Port Antonio, Dec. 1890, Hitchcock (MBG); Port Antonio, Aug. 1910, Lang 102 (ANSP); marsh near Black River, 12 Sept. 1907, Harris 9987 (NY); road- side, Hopeton, 13-22 Sept. 1907, Britton 1527 (NY); Och Bioa and vicinity, 4 April 190X, Britton ,fc Ilollick 2701, (NY); hanks of Cabaritta River, Meylerefield, 16 Dec. 1911, Harris 11808 (G, MBG, NY). Haiti: Prov. Barahona, Aug. 1910, Fuertcs 891 (NY); Prov. La Vega, July 1912, Fuertes 1768 (NY); Miragoanc and vicinity, July 1927, Eyerdam 201, 1,82 (G). Porto Rico: without locality, Read (ANSI'); Bayamon, 22 March 1885, Sintenis 1071, (MBG, NY); Aybonito, \i Nov. 1885, Sintenis 2010 (ANSP, P); Vega Baja, 5 Nov. 1013, Stevens akland, 25 July 1909, Mackenzie 4218 (MBG). Pennsylvania: without data, Miiehlenberg (Hot. Mus. Bcrl.-Dahl. type of V. rugosa, MBG phot.); Philadelphia, Nuttall (G); Whitcland, Chester Co., 26 June 1910, Bartram 1021 (G); vicinity of Conewago, 28 May L889, Small (US); York Furnace, 28 June 1899, MucKlwee 760 (MBG, NY); Mercersl.urg, Green (G). 1933] PERRY — NORTH AMERICAN SPECIES OF VERBENA 283 Maryland: Great Falls, 6 Aug. 1905, House 1863 (MBG); Hurlock, 29 May 1919, C. P. Smith 3186 (G). District of Columbia: Washington, 26 May 1889, Sudworih (G); Joy City, 19 June 1891, Blanchard (MBG). Virginia: Bluemont, 25 May 1905, House 875 (US); open woods near Churchview, 5 June 1921, Leonard & Killip 534 (US); north of Williamsburg, 17 June 1921, Grimes 3738 (NY); Bedford Co., June 1873, Curtiss (MBG); Chilhowie, 4 Aug. 1892, Small (F, NY); about Cumberland Gap, Lee Co., 27 July 1892, Small (F, G, MBG, US); Blue Ridge, 22 July 1891, Seymour 48 (G), 49 (MBG). North Carolina: near Chapel Hill, Ashe (US); Morgan town, 1872, Ruger (US); near Biltmore, 17 June 1S97, Biltmore Herbarium 4759b (G, NY, US); Madison Co., 27 July 1880, J. D. Smith (US). South Carolina: Abbeville District, July 1855, Hexamer & Maier (G). Georgia: Dalton, 9 Aug. 1900, Harper 385 (NY, US); Chickamauga Creek, near Ringgold, 6-12 Aug. 1895, Small (F, NY); between Taylor Ridge and Lafayette, 3 July 1900, P. Wilson 161 (NY, US). Florida: without locality, 1842-49, Rugel 127 (F, MBG, US). Alabama: Cullman, 29 May 1892, Mohr (US); Blount Springs, 5 May 1898, Baker (P); Blountsville, 6 May 1898, Baker (US); Choctaw Co., 13-15 Oct. 1896, Schuchert (US). Mississippi: Agricultural College, Oktibbeha Co., 11-17 Aug. 1896, Pollard 1312 (F, G, MBG, NY, P, US). Ohio: on limestone soil, Castalia, 27 July 1894, Moseley (G); Sandusky, 28 July 1894, Moseley (F, US); Oxford, 16 June 1910, Overholts (MBG). West Virginia: Shenandoah Junction, June 1891, Millspaugh 865 (NY); along Roanoke River, south of Roanoke, 2 June 1891, Small & Heller 432 (G); roadside, south of Williamsburg, 13 June 1921, Grimes 3716 (G); Sweet Springs, 14 Sept. 1903, Steele 814 (G, MBG, NY, US). Indiana: near boundary line to Galien, St. Joseph Co., 15 June 1911, Nieuwland 2685 (MBG, US); about 3 km. west of Goldsmith, 9 July 1913, Deam 13619 (MBG); about 1.6 km. west of Palmyra, 22 June 1916, Deam 20357 (G, US). Kentucky: without data, Short (NY); Stamping Ground, 29 May 1930, Singer 178 (US); Smithfield, 30 May 1909, Eggleston (NY); Nicholasville, 8 June 1923, McFarland 109 (MBG); Mammoth Cave, Edmonson Co., May 1899, Palmer (NY); Bowling Green, 21 May 1899, Price (MBG); near Franklin, 22 May 1926, Anderson 6 Woodson 47 (MBG); Kuttawa, 27 Sept.-9 Oct. 1909, Eggleston 5237 (MBG, NY). Tennessee: Carter Co., 8 July 1880, J. D. Smith (US); Knoxville, Ruth (F, P), 731 (US), 740 (NY), 765 (MBG); roadside at base of Chilhowee Mountains, Curtiss 1955 (F, G, MBG, NY); Chattanooga, 25 Aug. 1876, Engelmann (MBG); West Nash- ville, 26-27 May 1909, Eggleston 4430 (MBG, NY, US); Kingston Spring, 5 Aug. 1897, Eggert (MBG); Hollow Rock, Carroll Co., 14 Aug. 1897, Biltmore Herbarium 4759 (G, MBG, NY, US). Wisconsin: Baraboo, 1861, Hale (G, MBG). Illinois: Stony Island, June 1911, Greenman S64O (MBG); Stony Island, 25 June 1914, H. H. Smith 5923 (G, MBG); near 91st Street, Chicago, 28 July 1907, Greenman 1981 (MBG); South Chicago, 18 July 1913, H. H. Smith 5670 (G); Rock Island, 17 Aug. 1866, Engelmann (MBG); East St. Louis, 9 July 1898, Norton (MBG); bluffs near Prairie du Pont, 20 June 1876, Eggert (MBG); Red Bud, 3 June 1888, Pammel (MBG); Shawneetown, 26 May 1919, E. J. Palmer 15262 (MBG); Mound City, June 1859, Vasey (G). 284 ANNALS OF THE MISSOURI BOTANICAL (JARDEN I Vol. 20 Iowa: without locality, 17 July 1875, Arthur 24 (MHO); Cedar Falls, 12 July 1926, Pommel 272 ((!); Iowa City, Hitchcock (MBG); Columbus Junction, 27 Aug. 1899, Pammd 1699 (MBG). Missomii: Hannibal, 27 June 1 ( .*17, Davie 3588 (MBC); Ethel, 11 June 1915, Bush 7686a (MBC); Silcx, 29 May 1!)15, Davis 466% (MBG); prairie, St. Louis, May 1833, Engelmann 336 (MBG); banks of Mississippi above St. Louis, Oct. 1841, Engelmann (MBG); St. Louis, Sept. 1841, Engelmann (G, US); mineral region south- west of St. bonis, June 1845, King (MBC); Pacilic, 13 June 1897, Trelease 713 (MBG); Pacific, 3 June 1918, Oreenman 4'25 (MBG); Gray Summit, 16 June 1927, Kellogg 1 180 (M BG); Allcnton, 6 July 19! 1, LeUerman (MBG); near Sulphur Springs, 29 May 1927, Steyermark 292 (MBG); Crystal City, 14 May 1887, Trelease (MBG); Washington, 26 June 1888, Pammel (MBG); Victoria, 8 July 1890, Hitchcock (MBG); Jefferson Co., 20 June 1S76, Eggert (MBG); near Hillsboro, June, Riehl 466 (MBG); Hillsboro, 24 May 1886, F. Wisluenut 980 (MBG); near St. Genevieve, 27 May 1928, Oreenman \678 (MBC); Jerome, 24 May 1914, Kellogg 496 (MBG); Sheffield, 17 June 1916, Bush 761,7 (G, MBG, US); Westport, 17 May 1896, Bush 91/, (US); Independence, 1 June 1895, Tindall (MBG); south of Cedar Gap, 22 May-3 June 1911, Lansing 2976 (V, G); about 2 km. west of Mansfield, 5 12 June 1911, Lansing 3135 (F, G); Springfield, 31 July 1892, Dewart 42 (MBG); Wdlard, 24 July 1919, Blankituhip (P); limestone barrens, Ash (Jrove, 24 Aug. 1912, Standley 9336 (US); Carterville, 8 July 1910, /','. J. Palmer 8981 (MBG); Webb City, E. J. Palmer 200, 3319 (MBC); Swan, 24 Sept. 1899, Bush 450 (MBC); Galena, Stone Co., 27 May 1914, E. ./. Palmer 5769 (MBC); Gainesville, 26 June 1928, E. J. Palmer 34766 (MBG); Ea^le Bock, 24 June 1897, Bush 206 (MBG, IS); Noel, McDonald Co., 9 Sept. 1913, E. J. Palmer 4232 (MB< ',). ARKANSAS: Beaver Station, Kureka Springs, Olatfelter (MBG). Nebraska: Wahoo, June 1890, liydberg 154 (NY). Kansas: Lawrence, Stevens (US); between Heasanton and I'rescott, 20 June 1929, Rydberg A 1 »,ler 120 (MBG); sterile soil, Anderson Co., 1896, Hitchcock 791 (G, MBG, NY, US); Mound City, 18 July 18S7, Kcllermun (MBG). Ok i wioma: near Miami, 26 Aug. 1913, Stevens 2299 (G, N V); near Copan, 15 Aug. 1913, Stevens WHO (G); Koyil, 5 Aug. 1894, Bush 438 (MBC); on creek bank, near Pawhuska, 9 Aug. 1913, Steven* 1980 (G); Caddo, 20 June 1891, Sheldon 49 (US); near [dabel, IS May 1910, Houghton 3622, 3646 (G, MBG). This species appears to be very closely related to V. haslata. In its typical form, it is readily distinguished by its elongate spikes and narrow leaves. Often, however, intermediate forms occur between the two, making it somewhat difficult to find clear lines of demarcation between them. 18. V. Orcuttiana Perry, 18 n. sp. 18 V. Orcuttiana Perry, spec, nov., herbacea verisimiliter perennis; caulibus erectis quadrangulaiibua sparse pubeseentibus ramosis; foliis lanceolato-ellipticis spathulatisve in petiolum a latum attenuate 4 6 cm. longis grosse eerratis lineato- rugosis adpresao-pu bescentibus supra subtusque reticulatis dense patento-hirtellis; spicis pedunculatia compaetis elongatisque; bracteis lanceolato-acuminatis sparse ciliatis nervo medio et margine decurrentibus calyce brevioribus; calyce 4 mm. longo 1033] PERRY — NORTH AMERICAN SPECIES OF VERBENA 285 Stems several from a common base, 4-angled in cross-section, glabrous or very sparsely hirtellous, branching; leaves lanceolate- elliptical to spathulate, tapering into a margined petiole (1-2 cm. long), 4-6 cm. long, decussate, coarsely serrate, rugose and appressed-pubescent above, more densely spreading-pubescent or hirtellous beneath and prominently veined; spikes pedunculate, solitary or somewhat panicled, strict, mostly dense, elongate; rhachis more or less angulate; bracts lanceolate-acuminate, shorter than the calyx, midrib and margins more or less decurrent along the rhachis, sparsely ciliate; calyx 4 mm. long, appressed- pubescent and finely glandular, teeth short-subulate or acuminate, more or less connivent above the schizocarp; corolla-tube about as long as the calyx; corolla-limb 3-4 mm. broad; nutlets tri- gonous, 2 mm. long, raised-reticulate at the apex, striate toward the base; commissural faces extending to the tip of the nutlet, muriculate-scabrous. Distribution: Lower California. Specimens examined: Mexico: Lower California: Pinery, 27 July 1883, Orcutt (US); Hanson's Ranch, 29 July 1883, Orcutt (G, NY, US); table-lands, Hanson's Ranch, 30 July 1883, Orcutt 909 (G type); mountains, northern Lower California, 8 July 1885, Orcutt (US); Sierra Juarez, 12 July 1924, Gallcgos 2342 (US). This species, which has been passing as V. litoralis, is much like V. neomexicana var. xylopoda in the finely glandular spike and the angle of insertion of the flowers. It differs, however, in the shorter nutlets, the smaller corollas, and the type of the pubescence. In V. Orcuttiana, the trichomes are short and somewhat hirtellous and the pubescence of the inflorescence is closely appressed. In gross habit, it is scarcely to be distin- guished from V. simplex, but the latter has somewhat harsher pubescence and larger non-glandular flowers. Vent. Hort. Cels. 53, pi 53. 1800. Cav. Descr. 68. 1802. adpresso-pubescente tenuiter glanduloso; calycis dentibus breviter subulatis vel acuminatis; corollae tubo vix exserto; corollae limbo 3-4 mm. lato; coccis subtrigonis 2 mm. Iongis dorso striatis apice elevato-reticulatis; commissure muriculata. — Col- lected on table-lands, Hanson's Ranch, Lower California, 30 July 1883, Orcutt 909 (G), TYPE. 286 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 20 V. rigens Michx. FL Bor.-Am. 2:14. 1803. V. cuveifolia Raf. Med. Repos. N. Y. II. 5: 360. 1808. V. stricta ? mollis Torr. Ann. Lye. N. Y. 2: 234. 1827. V. mollis Haf. Atl. Jour. 146. 1832. V. strict a f. roseijlora ttenke, Rhodora 34: 10. 1932. V. stricta f. albijlora Wadmond, Rhodora 34: 19. 1932. Steins 3 12 dm. tall, subterete, simple or branched above, rather densely hirsute; leaves ovate or suborbicular, 6-10 cm. long, sessile or nearly so, sharply and mostly biserrate, thickish, hirsute and rugose above, densely hirsute- villous and prominently veined beneath; spikes solitary or several, short-pedunculate, thick, usually quite dense both in flower and in fruit; bracts lanceolate-subulate, approximately as long as the calyx, hirsute, ciliate; calyx 4-5 mm. long, densely hirsute, lobes acuminate; corolla protruding slightly beyond the calyx, pubescent without; corolla-limb 8-9 mm. broad; nutlets trigonous, 2.5 mm. long, raised-reticulate above, strongly striate below; commissural faces reaching tip of nutlet, muriculate or almost smooth. Distribution: eastern and central United Slates, from Pennsylvania westward through the Rocky Mountains. Probably introduced into other localities. Specimens examined: (Herb. Bot. Gard. Madrid type of V. Alopccurus, MUG phot.). Massachusetts: Fall River, 2 Sept. 1903, Hartford (G). Connectk t t: waste ground, Nangatuck, 19 July 1908, Blewitt 14 (G); Bridgeport, 27 Aug. 1892, Eame* (G). New York: pasture south of Pulpit Rock, Ithaca, 2 Aug. 1919, A. J. Fames 12797 (G). New Jiokskv: ballast, New Durham, 15 July 1S93, Van Sickle (US). Pennsylvania: along Lincoln Highway at Gap, Lancaster Co., Urban (G); vicinity of Conewago, Sept. 1892, Small (NY); Conewago, Sept. 1892, Heller 638 (US). Ohio: Dayton, Short (MBG). Michigan: Constantino, 2 July 192.'}, Fisher 19 (MBG); Pine Lake, Charlevoix Co., 18 Aug. 1917, Fhlers 642(F). Indiana: East Chicago, 10 Aug. 1910, Lansing 2810 (G); Lake Maximkuckee, 9 Aug. 1889, Evermann 970 (US); sand pit northeast of Winona Lake, 2 Aug. 1897, Beam (MBG); west of Palmyra, 22 June 1916, Deam 20857a (G). Kentucky: waste places, Hickman, 14 Aug. 1897, Biltmore Herbarium S653a (NY); Calvert City, Marshall Co., 10-19 June 1909, EggUston 4837 (NY); Wickliffe, 16 Aug. 1923, McFarland & Anderson 253 (MBG). Tennessee: Hickmann, Oatiinger (US); Henderson, June 1S92, Bain 828 (NY). Wisconsin: Mirror Lake, 15 July 1903, Eggert (MBG); Waupaca, 1907, Garesche (MBG); Trempeleau, 1861, Hale (G); Oregon road, Madison, 29 July 1889, Trelease (MBG). 1933] PERRY — NORTH AMERICAN SPECIES OF VERBENA 287 Illinois: Stony Island, Chicago, H. H. Smith 5645, 6028 (G, MBG), 5952 (G); Romeo, 26 July 1897, Umbach (MBG); Starved Rock, La Salle Co., June-Sept. 1921, Thone 88 (MBG); Oquawka, 1872, Patterson (G, NY); Bloomington, July 1886, Robinson (G); Carthage Township, 2 Sept. 1916, Gates 9994 (MBG); Decatur, 19 June 1896, Gleason 877 (G); Athens, Aug. 1863, Hall (MBG); East Hannibal, 13 June 1913, Davis 16 (MBG); Shepherd, 25 June 1915, Davis 6870 (MBG); Mississippi River, bluffs north of Alton, 4 Aug. 1910, Sherff (G); Bonpaa Township, 15 July 1925, Ridgway 2431 (MBG); E. St. Louis, 28 July 1900, Eggert (MBG); Grand Tower, 22 Aug. 1900, Gleason (G). Minnesota: Perham, Ottertail Co., 8 Aug. 1912, Chandonnet (MBG); St. Anthony, 7 July 1888, Schuette (G, NY); Willmar, July 1892, Frost (G); Morton, July 1890, MacMillan (P); Lake City, 28 July 1883, Manning (G); Winona Co., Aug. 1901, Holzinger (NY); Houston Co., July 1912, Freiberg (MBG). Iowa: Fayette Co., 12 July-4 Sept. 1904, Fink 251 (US); Hawkeye, Aug. 1896, Gardner 588 (NY); Cerro Gordo Co., 14 Aug. 1899, ./ones (MBG); Ames, 29 Aug. 1896, Pammel 85 (G, MBG, NY); Grinnell, Aug. 1877, Jones (NY, P); Mount Pleasant, 7 July 1898, Ball 1585 (MBG); Keosauqua, Aug. 1920, Graves 1947 (MBG); Ben- tonsport, July 1920, Graves 1994 (MBG). Missouri: suburbs of Hannibal, Davis 1202, 1518, 2791, 2960, 2962, 8227, 3589, 4461, 4487 (all MBG); Chain of Rocks, Aug. 1915, Beckwith 48 (MBG); St. Louis Co., 15 July 1872, Redfield 522 (MBG); St. Louis, Aug. 1838, Riehl 196 (MBG, NY); St. Louis, Aug. 1841, Engelmann (G); Allenton, Letterman (MBG); Williamsville, Wayne Co., 27 June 1914, E. J. Palmer 6107 (MBG); Jerome, 16 June 1914, Kellogg 498 (MBG); Warsaw to Linn Creek, Camden Co., July-Aug. 1913, Emig 107 (MBG); Cole Camp, Benton Co., 12 July 1897, Trelease 717 (MBG); railroad embankments, Mansfield, 5-12 June 1911, Lansing 3164 (G); Randolph, 17 July 1898, Mackenzie 261 (MBG); Independence, 26 June 1895, Bush 475 (MBG, NY); Willard, 9 July 1919, Blankinship (P); Carterville, 8 July 1910, E. J. Palmer 2980 (MBG); Webb City, 3 Aug. 1902, E. J. Palmer 199 (MBG); Turkey Creek, Joplin, 10 July 1897, Trelease 718 (MBG); Swan, 25 Sept. 1899, Bush 570 (MBG). Arkansas: waste places, Harvey 1958 (G, NY); Big Lake, 20-22 June 1911, McAlee 2058 (NY, US); Jonesboro, Craighead Co., 4 July 1927, Demaree 8553 (MBG); Fayetteville, Harvey 61 (MBG); Fort Smith, 1853-4, Bigelow (US). South Dakota: Big Stone Lake, Roberts Co., July 1922, Over 14386 (US); Wind- sor Township, Brookings Co., 27 July 1903, Johnson (MBG) ; Forestburg, 3 July 1910, Visher 4460 (MBG); 16 km. south of Interior, 29 June 1929, E. J. Palmer 37627 (G, MBG); near Fort Meade, Black Hills, 19 June 1887, Forwood 299 (US); canyons, Lead, 9 Aug. 1913, Carr 118 (G, MBG, NY, US); Rapid City, 22 July 1912, Visher 1507 (NY); Hot Springs, 3 Aug. 1892, Rydberg 982 (US). Nebraska: Lincoln, 27 June 1885, Webber (MBG); near Central City, 26 Aug. 1926, HeUer 14290 (MBG); South Fork of Platte, July 1856, H. Engelmann (G, MBG); Alma, 21 June 1897, Pammel (MBG); Anselmo, 8 July 1889, Webber (US); Broken Bow, 7 July 1897, Pammel (MBG); Callaway, 27 June 1901, Bates (G); Dismal River, south of Thedford, 27 June 1893, Rydberg 1422 (G, NY, US); North Platte, July 1896, Plank (NY); Ogallala, 16 June 1925, Jones (P). Kansas: Pottawatomie Co., 1895, Hitchcock 972 (US); prairie, Riley Co., 26 July 1895, Norton 891 (G, MBG, NY, US); Concordia, 24 July 1929, Benke 5164 (G, NY), forma roseiflora; hill 8 km. from Osborne, 11 July 1894, Shear 191 (G); Florence, 28-30 July 1903, Griffiths 5050 (US); vicinity of Caney, 29 June 1929, Rydberg & Imler 433 (MBG, NY); Syracuse, 28 July 1893, C. H. Thompson 154 (G, MBG, NY, US). [Vol. 20 288 ANNALS OF THE MISSOURI BOTANICAL GARDEN Oklahoma: Verdigris, 2 Aug. 1894, Bush 435 (MBG); 12 km. west of Pawnee, 27 July 1927, Slratton 2/,<) (MBC); near Tonkawa, 5 Aug. 1013, Stevens 1865 (G); Doby Springs, 20 Aug. 1027, Stratton 403 (MBG); near Shattuck, 11 Oct. 1913, Stevens 2080 (G); Norman, 10 Oct. 1914, Emig 365 (MBG); near Granite, 17 June 1913, Slrrms 1034 (G); Arbuckle Mountains near Davis, 23 June 1917, Emig 787 (MBG); vicinity of Fort Sill, 16 Aug. 1916, Clemens 11748 (MBG); near Cache, 25 June 1013, Strrens 1354V 2 (G, MBG); near Grant, 2 June 1916, Houghton 4000 (G, MBG, NY). Texas: Lipscomb, 1 July 1903, Howell 64 (IS); prairies, near Canadian, 11 Aug. 1900, lujucrt (MBG); Dallas, Reverchon (G, NY), 735 (MBC J). Montana: bottoms, Crow Agency, 14 July 1001, Blankinship (G). Wyoming: Whalen Canyon, 16 July 1804, A. Nelson 538 (G); Hartville, 20 July 1804, A. Nelson 505 (G, MBG). Colohado: Wray, 1-4 July 1919, Eggleston 15224 (MBG); Denver, 2 Sept. 1910, Eastwood 00 (G, MBG, US). New Mexico: creek bottom, Santa Fe, 1847, Femller 597 in part (ANSP, G). Washington: Meyers Falls, 20 Aug. 1902, Krcagcr 475 (O, NY, US). A very distinct species somewhat incapable of sharp delimita- tion on account of the tendency to hybridize with neighboring species. Normally it is recognized by its stout compact spike, imbricated flowers, and ovate-orbicular sessile leaves. 20. V. MacDougalii Heller, Bull. Torr. Hot. Club 26: 588. 1899. V. MacDougalii mut. rosella Cockerell, Am. Nat. 36: 809. 1902. Stems 3-8 dm. tall, stout, obtusely four-angled, simple or occasionally branched, cinereous-green, hirsute-pubescent; leaves oblong-elliptical or elongate-ovate, 0-10 cm. long, short-petiolate or narrowed into a subpetiolar base, coarsely and irregularly serrate-dentate, hirtellous, rugose and minutely pustulate above, densely pilose-pubescent and prominently veined beneath; spikes solitary or sometimes several, short-pedunculate, thick, comparatively dense both in flower and in fruit ; bracts lanceolate- subulate, for the most part noticeably longer than the calyx, pubescent, ciliate; calyx 4-5 mm. long, rather densely pubescent, lobes very obtuse, terminating in short subulate teeth; corolla- tube scarcely protruding beyond the calyx; corolla-limb 6 mm. broad; nutlets trigonous with convex back, 2.5 mm. long, raised- reticulate toward I he distal end, strongly or faintly striate below; commissural faces reaching tip of nutlet, muriculate or almost smooth. 1933] PERRY — NORTH AMERICAN SPECIES OF VERBENA 289 examined Wyoming Laramie Colorado: Palmer Lake, 22 July 1895, Osterhout (NY); La Veta, 14 July 1896, Shear 3577 (NY); Cucharas Valley, near La Veta, 20 July 1900, Vreeland 636 (NY); Stonewall, July 1912, Beckwith 138 (NY); Arboles, 10 July 1899, Baker 565 (F, G, MBG, NY, P, US). New Mexico: without locality, 1847, Fendler 597 in part (MBG); Vermejo Park, Colfax Co., 31 Aug. 1913, Wooton (NY); vicinity of Ute Park, Colfax Co., 2 Sept. Standley Standley Standley Read Ellis 258 (MBG, US); Santa Fe Co., 10 July 1847, Edwards (NY); Santa Fe Co., 1889, Brandegee (MBG); Santa Fe Co., 20 July 1898, Greene 77 (NY); Santa Fe Canyon, 3 Oct. 1913, Rose, Fitch & Parkhurst 17717 (US); Winsar's Ranch, Pecos River National Forest, 6 July 1908, Standley 4223 (F, G, MBG, NY, US); near Pecos, 15 Aug. 1908, Standley 4927 (F, G, MBG, NY, US); west of Las Vegas, St. Mulford mountains White P, US); Cloudcroft, 1912, Stearns 842, 358 (US); Cloudcroft, Aug. 1920, Schulz 255 (NY); Mescalero Reservation, Sacramento Mountains, 21 July 1905, Wooton (US). Arizona: Clark's Valley, 1 Aug. 1883, Rushy (NY, US); South Fork, Little Colorado River, Apache Forest, 23 Aug. 1920, Eggleston 17106 (NY); 16 km. south of Rowe's Point on the Grand Canyon, 5 Nov. 1899, Ward 11 (US); Flagstaff, 29 Aug. 1884, Jones (P); vicinity of Flagstaff, 8 July 1898, MacDougal 249 (ANSP, F, G, NY type); open pines, Flagstaff, 16 Aug. 1922, Hanson A148 (F, MBG). Utah: Mammoth Creek, near head of Sevier River, 10 Sept. 1894, Jones 6026 (MBG, NY, P). This southwestern representative of V. stricta is readily distin- guished by the short-petiolate elongated leaves, the compact spikes, and the floral bracts surpassing the calyx. 21. V. macrodonta Perry, 19 n. sp. PL 14. 19 V. macrodonta Perry, spec, nov., herbacea (basi ignota) verisimiliter perennis; caule 1-1.5 m. alto erecto hirsuto-hispidulo ramoso; foliis elongato-ovatis basi cuneata in brevem petiolum alatum attenuatis 10-14 cm. longis biserratis vel mu- cronulato-denticulatis supra scabro-hirsutis et inconspicue pustulatis subtus reticu- lars hirsutis; spicis paniculatis glanduloso-hirsutis basi foliolatis compactis deinde elongatis basi laxisque; bracteis lineari-lanceolatis subulatis ciliatis calyci subae- quantibus; calyce 5 mm. longo glanduloso aliquantulum viscido-pubescente; calycis dentibus 1.5 mm. longis subulatis; corollae tubo paulo exserto extus glabro vel puberulo; corollae limbo 5-6 mm. lato; coccis subtrigonis 2 mm. longis dorso sulcatis superiore parte tenuiter scrobiculatis; commissura muriculata. — Collected on the road from Miraflores to San Bernardo Ranch in Sierra La Laguna, Lower California, about 750 m. alt., 20 Jan. 1906, Nelson & Goldman 7425 (MBG), type. [Vol. 20 290 ANNALS OF THE MISSOURI HOTANICAL GARDEN Stem 1-1.5 m. tall, erect, branching, hirsute-hispidulous; ives elongate-ovate with cuneate base narrowed into a short -2 cm. long) margined petiole, 10-14 cm. long, coarsely and sharply biserrate-dentate with mucronate teeth, rugose with veins impressed and scabrous-hirsute above with minutely pustulate hairs, prominently veined and hirsute beneath; spikes panicled, subtended by leafy bracts, dense before anthesis, becoming elongated and open in fruit, glandular-hirsute; floral bracts lanceolate-linear, approximately equalling the fruiting calyx, subulate, ciliate; calyx 5 mm. long, glandular, somewhat viscid-pubescent, teeth 1.5 mm. long, subulate; corolla-tube protruding a little beyond the calyx, glabrous or puberulent without; corolla-limb 5-6 mm. broad; nutlets trigonous, 2 mm. long, shallowly scrobiculate on the upper half, tending to be sulcate toward the base; commissural faces muriculate. Distribution: Lower California. Specimens examined: MEXICO: Lower California: road from Miraflores to San Bernardo Ranch in Sierra La Lacuna, about 750 m. alt., 20 Jan. 1906, Nelson & Goldman 7^25 (MBG type, US). Verbena macrodonta is a coarse plant with large thickish leaves and open inflorescence, in a measure similar to V. MacDougalii, but differing in its less strict habit, somewhat remote fruits, more glandular calyces, and shorter plumper nutlets. 22. V. prostrata R. Br. in Ait. Hort. Kew. ed. 2, 4: 41. 1812. V. lasiostachys Link, Enum. Hort. Berol. 2: 122. 1822. Stems at first erect or ascending, at length spreading and ffusely branched, sparsely villous; leaves ovate or oblong-ovate th cuneate base tapering into a margined petiole, 5-10 cm. long, commonly dentate incised, veins impressed above, m both surfaces Dilose to soarselv v spikes solitary or more 4 often loosely paniculate, dense before anthesis, becoming elongated and open in fruit; bracts lanceolate- subulate, not longer than the calyx, villous and at times finely glandular; calyx 4-5 mm. long, villous or glandular-hirsute, subtruncate, the distal end connivent above the schizocarp, teeth subulate; corolla-tube a little longer than the calyx, very 1933] PERRY — NORTH AMERICAN SPECIES OF VERBENA 291 sparsely pubescent outside or apparently glabrous; corolla-limb 3-5 mm. broad; nutlets oblong-trigonous with convex back, 2-2.5 mm. long, slightly thicker at the distal end, raised-reticulate above, striate below, often with striae fading out toward the base; commissural faces more or less muricate, approximately reaching the tip of the nutlet. Distribution: Oregon and California. Specimens examined: Oregon: dry bank along railroad, about 1.5 km. north of Comstock, 18 June 1919, >son W Hammond 1249 above mouth of Rogue River, 8 July 1919, Peck 8702 (G, MBG, NY); in clearing about hotel, Agness, 22 June 1917, /. C. Nelson 1429 (G); Dryden, 16 June 1904, Piper 6160 (US); near Medford, Aug. 1922, Epling 5445 (MBG); south of Ashland, 19 May 1898, Applegale 2228 (US). California: near Yreka, Siskiyou, 19 June 1876, Greene 860 (F, G, MBG, NY); Yreka, 2 July 1910, Butler 1621 (MBG, P, US); Mount Shasta, 13-27 July 1892, Palmer 2529a (US); Soda Springs, Nevada Co., 30 July 1881, Jones 2598 (P); Round Valley, Mendocino Co., 20 May-20 June 1898, Chestnut (US); near Clear Lake, 1865, Torrey 417 (G, NY); Clear Lake, Lake Co., 12 July 1929, Blankinship (MBG); southern slope of Mount Sanhedrin, Lake Co., 19 July 1902, Heller 5919 (G, MBG, NY, P, US); west of Windsor, near Russian River, Sonoma Co., 27 June 1902, Heller 5785 (F, G, MBG, NY, P, US); Oak Knoll, 5 May 1901, Braunton 378 (US); Irish- town, Amador Co., June 1893, Hansen 964 (MBG); West Point Bridge, Calaveras Co., 7 July 1896, Hansen 1823 (US); Stanford University, Santa Clara Co., April 1898, Abrams (P); San Jose, 14 July 1899, Pammel 187 (MBG); Santa Cruz, 21 June 1881, Jones (G, P); in pine woods, Pacific Grove, 25 May 1903, Heller 6778 (F, G, MBG, NY, P, US); Carmel Bay, Sept. 1902, Elmer 404B (G, P); Monterey, Hartweg 1924 (G, NY); Monterey, Aug. 1917, Parish 11590 (P); Santa Lucia Mountains, May 1898, Plaskett 142 (NY, US); San Antonio River, 1880, Vasey (US); Wood's Creek, Fresno Co., 19 June 1910, Clemens (P); San Luis Obispo, 26 June 1876, Palmer 342 (MBG, NY, US); San Luis Obispo Co., 19 June 1887, Summers (MBG, P); Santa Barbara Co., 1865, Torrey 416 (G, NY); Santa Barbara, May 1902, Elmer 3846 (MBG, P, US); seep, base of ocean bluffs, 32 km. northwest of Santa Barbara, 26 March 1925, Munz 9295 (P); Sulphur Mountain Spring, Ventura Co., 1-2 June 1908, Abrams & McGregor 5 (G, NY, US); Ventura, 13 April 1923, Kendall (P); San Buenaventura, March 1861, Brewer 229 (US); Los Angeles, 1860-62, Brewer 31 (G, US); Los Angeles, 29 May 1891, Fritchey 29 (MBG); near Mesmer, 10 June 1917, Johnston 1320 (P, US); near Pasadena, June 1893, Haynes (P); San Antonio Canyon, 28 June 1917, Johnston 1608 (P); edge of woods, Palomar Mountain, 11 Aug. 1918, Spencer 994 (G, P); wayside, Mesa Grande, 2 June 1919, Spencer 1164 (G); Spencer Valley, near Julian, San Diego Co., 20 June 1903, Abrams 3787 (F, G, MBG, NY, P); bottom of canyon south of "Lemon Tank," west coast, San Clemente Island, 10 April 1923, Munz 6734 (P). Although, from the original publication, one would naturally 292 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 20 infer that V. prostrata is Aiton's species, it is in all probability Robert Brown's. Aiton, in his acknowledgments (postscript to the fifth volume of Hortus Kewensis), mentions the new matter added by his friend Robert Brown, some without reference to his name; more tangible evidence is furnished by Schauer, who, in his monograph, indicates the specimen at Kew as V. prostrata R. Br. The species is comparatively easy to distinguish by its de- cumbent habit, soft villous pubescence, and elongated spikes. It closely resembles V. robusta, which is much harsher and of limited distribution. The following specimens differ from the species in having scarcely glandular inflorescence and fruiting calyx only 2.5- 3 mm. long shortly conniving beyond the fruit: Califoknia: Three Rivers, Tulare Co., 9 July 1904, Culberlson 4210 (MBG, P); San Bernardino, Sept. 1886, Parish 8. Stems (i-9 dm. high, erect, paniculately branched above, glabra te or sparsely hirsute; leaves ovate or oblong-ovate with cuneate base tapering into a margined petiole, 4-7 (-10) cm. long, usually 3-cleft, irregularly serrate-dentate with apiculate teeth, or incised, rugose and scabrous-pubescent above, less harshly pubescent beneath with veins prominently reticulate; spikes often crowded, subsessile, usually dense but occasionally elongated; bracts lanceolate-subulate, more or less exceeding the calyx in length, densely glandular-hirtellous; calyx 4 mm. long, densely glandular-hirsute, lobes obtusish, terminating in very unequal acuminate-subulate teeth; corolla-tube a little longer than the calyx, puberulent without; corolla-limb 3-4 mm. broad; nutlets oblong-trigonous, 2-2.5 mm. long, raised-reticulate above with striae fading out toward the base; commissural faces more or less muricate, approximately extending to the tip of the nutlet. Distribution : California and Lower California. Specimens examined: 1933] PERRY — NORTH AMERICAN SPECIES OF VERBENA 293 California: vicinity of lone, Dec. 1904, Braunton 1268 (MBG, NY); Tiburon, Marin Co., 26 July 1900, Eastwood (G); valley back of Berkeley, 23 Oct. 1880, Engelmann (MBG); West Berkeley, May 1887, Greene (US); salt marshes, Oakland, Sept. 1886, Congdon (G); Temescal, Alameda Co., Aug. 1891, Michener & Bioletti 128 (G); Crystal Springs Lake, San Mateo Co., July 1903, Elmer 4950 (MBG, P, US); San Luis Obispo, 1876, Palmer 841 V 2 (F, G, NY, US); west Sherman Canyon, 18 June 1901, Braunton 99 (US); Valdez Bay, Santa Cruz Island, 5 Sept. 1927, Jones (P); sandy soil near coast, Santa Catalina Island, 19 July 1915, Macbride & Payson 850 (G); Santa Catalina Island, 20-25 July 1917, Eastwood 6500 (US); dry stream- bed, pebbly beach canyon, Santa Catalina Island, 13 May 1928, Dunkle 1955 (P); Avalon, Santa Catalina Island, Sept. 1896, Trask (MBG); Avalon, May 1897, Trask (F, US); Avalon, Aug. 1901, Trask (NY); Avalon, 28 April 1914, Carlson (US); Avalon, 13 June 1915, Carlson (G, MBG); Jamuel Valley, San Diego Co., 26 June 1875, Palmer (G), 310 (F, MBG, NY). Mexico: Lower California: near Rancho Salina, foot of Guatay Grade, about 6 km. south of Rio Guadalupe, 11 Sept. 1929, Wiggins & Gillespie S977 (MBG); Santa Tomas, 15 July 1885, Orcutt 1801 (MBG, US). Verbena robusta has been much confused with the nearly- related V. prostrata, but is quite readily distinguished from the latter by the brighter green color of the herbage, the scabrous surface of the leaves, and More over, the mature calyx lacks the marked tendency toward sub- connivent lobes, a characteristic of V. prostrata. The schizo- carps of the two are very much alike. The collection Orcutt 1301 has a greatly elongated inflorescence and the leaves are not particularly scabrous. The specimens Braunton 99 and 1263 show unusually long floral bracts. These are probably atypical phases of the species or possibly hybrids. 24. V. xutha Lehm. Del. Sem. Hort. Hamb. 7, 8. 1834; Linnaea 10: Litt.-Ber. 115. 1835-6. V. strigosa Hook. Comp. Bot. Mag. 1: 176. 1836, not V. strigosa Cham. Linnaea 7: 256. 1832. V. Lucaeana Walpers, Rep. 4: 23. 1844-48. Stems upright, 6-10 dm. tall, branched, hirsute-hispid; leaves oblong or broadly ovate in outline, 5-8 (-12) cm. long, incised- pinnatifid or more often trifid with segments coarsely dentate, the lateral much smaller than the middle segment and close to the sessile base of the blade, strigose above, canescent and spread- ing-hirsute below, trichomes particularly prominent along the somewhat paler midrib and veins; spikes elongate, somewhat strict, not compact except at an thesis, short-pedunculate; bracts 294 ANNALS OF THK MISSOURI liOTANICAL (SAKDKN [Vol. 20 lanceolate-subulate, subequalling or commonly a little shorter than the calyx, strigillose, ciliate; calyx 3-4 nun. long, strigose- hirsute, lobes acuminate-subulate; corolla-tube approximately as long as the calyx, the throat pubescent without; corolla-limb 5-8 mm. broad; nutlets 2 mm. long, raised-reticulate on the upper half, faintly striate below, commissural faces reaching the apex of the nutlet, inuricate or muriculately scabrous. Distribution: Alabama to Texas. Specimens examined: Alabama: Navy Cove, Aug. 1889, Mohr (US). Louisiana: without data, Hale (G, MBG), Drummond (G); vicinity of Alexandria, 9 June 1899, Ball 605 (MBG, US); Chopin, 6 May 1915, E. J. Palmer 7556 (MUG); St. Martinville, 30 May 1893, Langlois (MBG, US); Pointe a la Hache, 25 June 18X1, LangUris 123 (US); New Orleans, 1832, Drummond 253 bis (K type of V. strigosa); near Schriever, 8 June 1917, Munz 1607 (P); Cameron, 4 July 1903, Tracy 8708 (F, G, MBG, NY, US). Arkansas: near Homan, 10 June 189X, Eggert (MBG); Tcxarkana, Aug. 1881, Lctterman (MBG, US). Texas: Gladewater, 18 June 1901, Reverchon 2532 (MBG, NY); near Longview, 7 June 1S99, Eggcrt (MBG); Corsieana, 3 Oct 1900, Reverchon 2118 (MBG, US); Huntsville, 1920, Warner 43 (US); sandy land, Brazos Co., 1S99, Ness (G); Brazos, July 1S99, IAndheimer (MBG); dry banks of Brazos River, 23 June 1917, Munz ItfO (P); College Station, 24 July 1899, Reverchon (M IK J); black land prairie, Montgomery Co., 18 21 July 1900, Dixon 473 (F, G, NY); Burton, 26 May 1872, Hall 434 (F, G, MBG, NY, P, US); Austin, 12 July 1920, Tharp 667, 668 (US); Austin, Aug. 1921, Sckulz 67.9, 701 (US); San Marcos and vicinity, 1898, Stanfield (NY); Industry, 1844, IAndheimer 146 (MBG); Cypress City, May 1877, /. Ball (MBG); near Houston, April 1842, IAndheimer 154 (G, MBG); swamps, Houston, Aug. 1904, Kuntze 23811 (NY); Lotus, about 16 km. west of Houston, 8 Aug. 1921, Ferris & Duncan 3268 (MBG); Strand, Jefferson Co., 9 April 1924, Tharp 3166 (MBG); Galveston, 6 June 1920, Fisher 212 (US); Columbia, 5 Oct. 1900, Bush 1275 (MBG); Columbia, 23 Sept. 1901, Hush 899 (MBG); Ilallettsville, 9 Aug. 1912, Fisher 122 (US); Ottim 30 Aug. 1926, Bogusch 1235 in part (P); Sequin, 17 June 1903, Groth 187 (F, G, NY, US); Cave Lake, Jackson Co., 30 June 1915, Druehd 2842 (M BG). This species is easily distinguished by its erect habit, coarse pubescence, elongated open spike, and sessile commonly trifid leaves. It has often been confused with V. canesceyis var. RoewHTiana, but the latter is a smaller and more compact plant. 25. V. olicata G 1903. Coarse herb with stems decumbent to ascending, branched, hirtellous; lower leaf-blades elliptic-ovate, narrowed into a d the same length, 1-3 (-4) long, broadly obtuse, plicate, coarsely incised-dentate, often 1933] PERRY NORTH AMERICAN SPECIES OF VERBENA 295 3-lobed, more or less canescent, rugose and somewhat appressed- hirsute above, hirsute beneath and prominently marked (particu- larly near the margin) with whitish veins; upper leaves similar but smaller, often appearing spathulate; spikes terminal on stems and branches, ordinarily not compact; bracts ovate- lanceolate, usually exceeding the calyx, at times barely equalling it, acute, hirsute, midrib often noticeable; calyx 3.5^4 mm. long, more or less glandular-hirsute, lobes very obtuse or subtruncate, terminating abruptly in subulate teeth; corolla-tube scarcely longer than the calyx; corolla-limb 4-6 mm. broad, anterior lobe retuse; nutlets cylindric-trigonous, 2-2.5 mm. long, shallowly scrobiculate above, changing to indefinitely striate toward the base; commissural faces finely muricate-scabrous to practically smooth, not reaching the tip of the nutlet. Distribution: Texas, New Mexico, Arizona, and northern Mexico. Specimens examined: Texas: without data, Berlandier 2506 in part, 644 (MBG); without locality, Sept. 1881, Havard (US); southwestern Texas, Reverchon 118 (G); western Texas, 1851-2, Wright 1496 (G, MBG, NY); sandy roadside, Chillicothe, 28 Sept.-3 Oct. 1906, Ball 1171 (US); sands, Estelline, 25 May 1904, Reverchon 4314 (MBG); sandy waste, Garza Co., 6 June 1925, Ruth 1289 (US); Post, 22 May 1925, Wooton (US); Lubbock, 24 April 1930, Demaree 7589 (US); moist open ground along creeks, Sweetwater, Nolan Co., 27 May 1918, E. J. Palmer 18780 (MBG); north of Colorado, Mitchell Co., June 1900, Eggert (G, MBG); prairie north of Stanton, Martin Co., 13 June 1900, Eggert (MBG); rocky and sandy soils, Comanche, 8 Aug. 1877, Reverchon 834 in part (MBG); near Comanche, 10 May 1900, Eggert (G, MBG); rocky prairies, Brown Co., 10 Aug. 1877, Reverchon 787 in part (MBG); Brown Co., April 1882, Reverchon 787 (US); San Angelo, 19 May 1903, Reverchon 1958 (MBG); Barstow, 14 April 1902, Tracy & Earle 80 (F, G, MBG, NY), type collection; Barstow, 15 April- 3 May 1902, Tracy & Earle 41 (NY); Oxona, 13 April 1930, Jones 26221 (P); Davis Mountains, 5 Aug. 1918, Young (P); cliffs back of Fort Davis, Davis Mountains, 9-12 July 1921, Ferns & Duncan 2726 (MBG, NY, P); about 5 km. east of Study Butte, Brewster Co., 30 June 1931, Moore & Steyermark 8299 (MBG); Alpine, 7 June 1926, E. J. Palmer 80523 (MBG); near Boquillas, 17 April 1919, Hanson 619 (MBG, US); Austin, 12 May 1872, Hall 429 (F, G, NY); gravel bars of Blanco River, Blanco, 5 April 1918, E. J. Palmer 13282 (MBG); near Feodora, Terrell Co., 26 April 1928, E. J. Palmer 33537 (MBG, NY); Devils River, Valverde Co., May 1913, Orcutt 6235 (MBG); Corpus Christi Bay, Dec. 1879, Palmer 2038 (G); Laredo, Aug. 1879, Palmer 2040 (G); Laredo, Berlandier 1485 ( = 225) in part (G); Laredo, 21 March 1903, Reverchon 3904 (G, MBG, US); Laredo, 1913, Orcutt 5555, 5717 (MBG); near Pharr, Hidalgo Co., 6 April 1931, McKelvey 1756 (G). New Mexico: plains, Carrizoza, 8-19 May 1902, Earle 606 (NY); in valley near Gray, 26 July 1900, Earle 427 (NY, US); neighborhood of San Miguel, 12 Aug. 1847, Fendler 594 (G, MBG). Arizona: desert prairie, north of Tucson, 24 April 1913, Greenman & Greenman 28 296 ANNALS OF THE [Vol. 20 (MBG); Tucson, 2 May 1892, Tourney S06 (US); slopes west of Tucson, 30 Dec. 1010, Bartram 256 (ANSP); Cienega, near Pantano, 14 June 1881, Pringle (F, G, MBG); valley near C:unp Lowell, 8 June 1882, Pringle (ANSP, F, NY); near Fort Lowell, 15 Sept 1900, Griffiths 1595 (NY). Mexico: Chihuahua: Colonia Juarez, Sierra Madre Mountains, 11 Sept. 1003, Jones (P). COAHtXILA: Sierra Mojada Mountains, 10 April 1802, Jones 372 (P, US). In habit V. plicaia is somewhat similar to V. neomexicana, but in inflorescence it strongly resembles V. canescens var. Roemeriana. Its distinctive character is foliar and is most easily seen in the basal and the lower stem-leaves. The leaf is obviously petioled, often 3-lobed, and very shallowly incised-dentate; moreover, the veins beneath are whitish and particularly prominent near the margin owing to the plication of the leaf and the apparent broadening of the veins in this region. neomexicana (( iray) Small, Fl. Southeast. U. S. ed 1010. 19(K*. anded Gray, Syn. Fl. N. Am 1878. V. officinalis var. hirsute Torr. Hot. Mex. Bound. 2: 128. 1859. Plant slender; stems upright, branched, hirsute; leaves 1-5 cm. long, pinnately cleft or almost parted, segments incised or coarsely toothed, rugose, somewhat scabrous and finely pustulate above, the veins more or less prominent beneath, hirsute on both sur- faces; spikes solitary or tending to be panicled, usually short- pedum- led, hirsute; bracts lanceolate-acuminate, commonly not longer than the calyx; calyx about 3 mm. long, hirsute-pubescent and very slightly glandular, teeth short and subulate; corolla- tube scarcely longer than the calyx; corolla-limb approximately 4 mm. broad; nutlets trigonous with convex back, 2 mm. long, very shallowly retieulate-scrobiculate on the upper half, longi- tudinally striate below; commissural faces extending to the tip of the nutlets. Distribution: Texas and New Mexico. Specimens examined: Texah: Fort Davis, 13 Sept. 1018, Young 1 70S (US). Nrcw Mexico: White Mountains, Lincoln Co., 12 Aug. 1807, Wooton 6^6 (NY); Ruidoso Creek, Lincoln Co., 3 July 1805, Wooton (NY, US); Kingston, Sierra Co., 6 July 1904, Metcalfe 955 (US); borders of thickets, near Coppermines, 1851, Wright 1A97 (G typk, MBG, NY. US); Pinos Altos Mountains. 1880. Greene (F. MUG): 1933] PERRY — NORTH AMERICAN SPECIES OF VERBENA 297 bottom of Tierra Blanca Canyon, Gila Forest, 18 Aug. 1916, Chapline 609 (NY); G. 0. S. Ranch, vicinity of Silver City, 27 Aug.-12 Sept. 1911, Holzinger (US); Mogollon Mountains, on or near West Fork of Gila River, Socorro Co., 28 Aug. 1903, Metcalfe 612 (G, MBG, NY, P, US). The collections of Young 1708 , Wooton 646, and Wooton (collection of July 3, 1895) are by no means typical of the species, but perhaps are better placed here than elsewhere. This species appears to be very closely related to V . canescens and V. gracilis. It differs from both, however, in the upright habit and the nutlets. In both V. canescens and V. gracilis the commissural face does not extend to the tip of the nutlet and the striae on the dorsal surface are less conspicuous. 26a. Var. xylopoda Perry, 20 n. var. Stem somewhat coarser; pubescence shorter, denser and more glandular; calyx 4 mm. long, glandular-hirsute, teeth acuminate; corolla-limb 6-10 mm. broad. Distribution: Arizona, California, and northern Mexico. Specimens examined: Arizona: Clarkdale, 17 Sept. 1921, W. W. Jones 344, 846 (G); Rio Verde, 8 Sept. 1865, Coues & Palmer 571 (MBG); Skull Valley, 28 April 1903, Jones (MBG, P, US); near Oracle, 20 April 1930, Harrison & Kearney 6689 (NY, P); Chiricahua Moun- tains, Cochise Co., 1 May 1894, W. W. Price (P); Paradise, Chiricahua Mountains, 16 Sept. 1907, Blumer 2170 (F, US); Chiricahua Mine, 21 Oct. 1907, Blunter 1804 (F, G, MBG, NY, US); Warren, Cochise Co., 20 May 1915, Carlson (US); Bisbee, 3 Oct. 1892, Mearns WIS (US); near Fort Huachuca, 1882, Lemmon 2857 (G); Fort Huachuca, Aug. 1892, Wilcox (NY); Huachuca Mountains, 3 Sept. 1903, Jones (P); Ash Canyon, Huachuca Mountains, 6 Aug. 1909, Goodding 834 (G, NY); Huachuca Mountains, 3 Sept. 1928, Harrison & Kearney 5796 (US); Santa Catalina Mountains, 27 July 1917, Munz 1149 (P); Sabino Canyon, Santa Catalina Mountains, 21 April 1922, Hanson A 1130 (MBG type); about 13 km. south of Vail, 31 Aug. 1903, Jones (P); foothills of the Santa Rita Mountains, 11 May 1884, Pringle (ANSP, F, G, US); Santa Pdta Forest Reserve, 8-13 Sept. 1902, Griffiths 3431 (US); Santa Rita Range Reserve, 12 May 1912, Wooton (US); slopes about Calabasas, 21 April 1908, Tides- trom 872 (US); Baboquivari Mountains, 28 March 1927, Peebles, Harrison & Kearney 3790 (US); Baboquivari Mountains, 26 Sept. 1927, Harrison 4778 (US); Baboquivari Mountains, 12 April 1928, Gilman 1120 (P). California: without locality, 1876, Palmer (G), SS9y 2 (US). Mexico: Lower California: Big Canyon of Tantillas Mountains, 10 Sept. 1875, Palmer (G), S12 (F, MBG). 20 Var. xylopoda Perry, var. nov., typicam simulans sed indumentum brevius densiusque; calyce 4 mm. longo gland uloso-hirsu to; calycis dentibus acuminatis; corollae limbo 6-10 mm. lato. — Collected on rocky slopes, Sabino Canyon, Santa Catalina Mountains, Arizona, 21 April 1922, Hanson A11S0 (MBG), type. [Vol. 20 298 ANNALS OF THE MISSOURI BOTANICAL GARDEN Sonoka: Guadalupe Canyon, 27 Aug. 189:5, Merton 2042 (US); "Niggerhead Mountains, near monument no. 82," Aug. 1893, M earns 1887, 1918 (US); Fronteras, June 1851, Thurher 446 (G, NY). Chihuahua: Saint Eulalia Hills, 30 July 1885, Wilkinson (US); San Diego, 26 April 1S91 , Hart man 608 (G); rocky hills, near Chihuahua, 16 April 1885, Pringle 270 (ANSP, P, G, US); vicinity of Chihuahua, 8-27 April 1908, Palmer SB (F, G, MBG, NY, US); "between Sacramento and Chihuahua," 24 Aug. 1846, Wislizenns 150 (MBG); in the Sierra Madre, 21 June-29 July 1899, Nelson 6161 (G, US); near Colonia Garcia, 29 July 1899, Townsend & Barber 192 (F, G, MBG, NY, P, US); between Colonia Garcia and Pratt's Ranch, below Pacheco, 22-24 Aug. 1899, Nelson 6871 (G, US). This variety differs from the species in the denser and more glandular pubescence and the larger corolla. The nutlets also are slightly longer, with the reticulations somewhat deeper than in the species, and the commissural faces hardly extend to the tip of (he nutlets. The specimens Nelson 6161, 6271 and Town- send (St Barber 102 closely resemble the above variety in in- florescence, but are more like V. neomexicana in the long and somewhat sparsely hirsute indument on the lower part of the stem; Pringle 270 and Wilkinson approach variety hirtella. 26b, Var. hirtella Perry, 21 n. var. Plants densely canescent-hirtellous; leaves more or less shal- lowly incised; bracts usually broadly ovate-acuminate; corolla- limb about S mm, broad. Distribution: Texas, New Mexico, and Coahuila, Mexico. Specimens examined: Texas: rocky places between Van Horn Wells and Muerte, 2 July 1852, Parry, Bigelow, WrigtU & Schott (NY, US); Sivermore Peak, Davis Mountains, 9-12 July 1921, Ferris d' Duncan 2607 (MBQ, NY); sand bars of creeks, Davis Mountains, 11 July 1920, E. ./. Palmer $0791 (MBG); Valentine, 28 April L930, Jones 26224 in part (P); near Shnfler, Presidio Co., 26 April 1931, McKelvey 2046 (G); Pinto Canyon, near Ruidosa, 13 April 1919, Hanson 646 (G, NY, US); Chisos Mountains, 12 Aug. 1916, Young 112 (MUG); foothills of Chisos Mountains, 22 May 1928, E. J. Palmer 84065 (MUG type, NY); gravelly mesa, north side of Chisos Mountains, 27 June 1931, Moore & Steyermark 8277 (MBG). New Mexico: Socorro Mountains, 11 July 1897, Herrick 715 (US); low moun- tains west of San Antonio, 14 April 1908, Wooltm 8852 (US). Mexico: Coahuila: Sierra de Parras, Purjrus 1094 (F f G, MBG, NY, P). The pubescence of this variety is much finer and shorter than 21 Var. hirtella Perry, var. nov., planta dense liirtello-caneseens; foliis plus minusve breviter incisis; bracteis fere late ovato-acuminatis; coiollae limbo circiter 8 mm. lato. — Collected on the foothills of the Chisos Mountains, Texas, 22 May 1928, E. J. Palmer SJfi6B (MBG), type. 1933] PERRY — NORTH AMERICAN SPECIES OF VERBENA 299 in the above. The leaves are not so deeply incised and often tend to be elongated. The leaves of Purpus 1094 are so narrow and shallowly incised that it appears superficially like V. perennis; nevertheless, the character of the pubescence allies it with this variety. 27. V. perennis Wooton, Bull. Torr. Bot. Club 25 : 262. 1898. Stems several from a woody base, divaricately ascending- erect, more or less strictly branched, glabrate or often finely glandular and slightly hispidulous with short stiff antrorse hairs; leaves predominantly linear, 1-4 cm. long, entire or pinnately few-lobed, erect-ascending, sparsely hispidulous, mar- gins revolute; spikes terminal, pedunculate, slender-filiform, elongate; bracts ovate, 1.5-3 mm. long, acute, hispidulous, ciliate; calyx 4-5 mm. long, pubescence more abundant along the nerves, lobes subequal, short, acute; corolla-tube slightly longer than the calyx, pubescent; corolla-limb 5-7 mm. broad, lobes repand ; fruit more or less remote ; schizocarp 3 mm. long, strongly constricted along the lines of cleavage; nutlets subcylindric, reticulate-scrobiculate except at base; commissural faces smooth or slightly scabrous and not extending to the tip of the nutlet. Distribution: Texas and New Mexico. Specimens examined: Texas: Glass Mountains, 25 Aug. 1925, Tharp 3682 (US); Loyola, 5 Nov. 1902, Seler (G); Guadalupe Mountains, 1882, Havard 197 (G); Guadalupe Mountains, 13 Aug. 1916, Young (MBG); rough grassy slopes, McKittrick Canyon, Guadalupe Mountains, 23 July 1931, Moore & Steyermark 3611 (MBG); Guadalupe Mountains, 8 Aug. 1931, Clarke 4250 (MBG). Queen, Aug. 1909, Wooton Standley Standby (US); White Mountains, Lincoln Co., 21 July 1897, Wooton 187 (G, MBG, NY, P), type collection; Ruidoso Creek, White Mountains, 5 Aug. 1901, Wooton (MBG); north of El Capitan Mountains, Lincoln Co., 31 Aug. 1900, Earle 387 (MBG, NY, P, US); Capitan, 8-19 May 1902, Earle 619 (NY); Gray, Lincoln Co., 6 June 1898, Skehan 20 (F, G, MBG, NY, P, US); Berendo Creek, 13 May 1905, Metcalfe 1568 (F, G, MBG, NY, US). The relationship of this species is somewhat anomalous. The lobing of the leaves and the character of the nutlets seem to ally it with V whereas and the predominance of practically entire linear-oblong leaves recall V. simplex. It could scarcely be confused with either [Vol. 20 300 ANNALS OF THE MISSOURI BOTANICAL GARDEN since the character combination of an open spike, very narrow leaves, and sparsely short-strigillose hairs is not found elsewhere in the group under consideration. 28. V. gracilis Desf. Cat. Hort. Paris, ed. 3, 393. 1829. V. remote Benth. PL Hartw. 21. 1839. V. arizonica Gray, Proc. Am. Acad. 19: 95. 1883. Low diffusely branched herb; branches decumbent to ascending, more or less glandular, canescent, hirsute; leaves ovate, cuneate at base and narrowed into a margined petiole, 1-3 cm. long, incised-pinnatifid to | (innately cleft, segments oblong, subincised, acutish, midrib and veins impressed, sometimes glandular, hirsute particularly on the lower surface; spike terminal, slender, more or less sparsely flowered, sessile, the lowermost flowers often appearing in the axils of the upper leaves; bracts linear, attenuate, gradually reduced in length toward the tip of the spike, usually much longer than the calyx; calyx 3 mm. long, more or less glandular, hirtellous, subtruneate, teeth minute and subulate; corolla inconspicuous, the tube somewhat longer than the calyx, the limb about 2-3 mm. broad; nutlets 1.5-2 mm. long, finely scrobiculate from the apex practically to the base; commissural faces muricately scabrous. Distribution: Arizona, Utah, and Mexico. Specimens examined: (Par. type, MBG phot). Described from specimen cultivated in the Botanical Garden of Paris. Arizona: Gardiner's Spring, 25 June 1882, Pringle (ANSP, F, G, NY); Gulching Ground, Pine Canyon, Chiricahua Mountains, 5 July 1907, Blumer 1612 (F, G, MBG, NY, US); San Pedro River, Mexican Boundary Line, 12 Oct. 1892, Mearns 1116 (US); Tanner's Canon, near Fort Huachuca, 1SS2, Lemmon (G); Fort Huachuca, 1890, Patzky (IS); Fort Huachuca, Wilcox (NY, US), 29 (US). Utah: mesa cast of Monticello, 25 July 1911, Hydberg & Garrett 9201 (NY). Mexico: Chihuahua: waste places, Chihuahua, 26 May 1885, Pringle 64 (G); near Chihua- hua, 7 May 1887, Pringle (F, MBG); vicinity of Chihuahua, 1-21 May 1908, Palmer 200 (US). Duran:> (US); Fort Meade, March 1880, /. D. Smith (VS); between Tallahassee and St. Marks, April 1843, Hugcl (NY); sandy open ground near Tallahassee, 9 April 1929, E. J. Palmer 35235 (G, MBG); Aspalagii, Chapman (MBG); dry pine barrens, near Apalachicola, 11 Aug. 1889, Biltmorc Herbarium 4761a (G, NY, US). Alabama: without locality, 1S59, Beaumont (G); Auburn, bee Co., 10 May 1890, Earle & Underwood (NY); Auburn, 10 Feb. 1897, Earle narentlv inserted in a depression ted lines of cleavage; connective of the upper anthers chiefly appen- daged. Herbaceous perennials with prostrate, decumbent, as- cending, and sometimes erect stems. Flowers usually showy, at first fascicled or somewhat corymbose, later spicate. Calyx usually more than twice as long as the schizocarp and constricted or contorted beyond it. Species 33-51. 19331 PERRY — NORTH AMERICAN SPECIES OF VERBENA 311 Key to the Species A. Leaves tapering into a margined petiole or subsessile or sessile, at least not subauriculate and semiamplexicaul at the base. B. Nutlets with a very definite beak parallel to the axis of the schizocarp. . 33. V. quadrangulata B. Nutlets with an indefinite beak or none. C. Nutlets suggesting a tendency toward a beak (a slight protrusion horizontal to the axis of the schizocarp); commissural face practi- cally reaching the tip of the nutlet 34- V. delticola C. Nutlets without a beak; commissural face not reaching the tip of the nutlet. D. Nutlets subovoid, lateral surfaces ventricose and smooth, not at all similar to the dorsal surface 39. V. tumidula D. Nutlets subcylindrical, lateral surfaces not ventricose and usually scrobiculate, similar to the dorsal surface. E. Leaves shallowly lobed, incised or toothed. F. Corolla-tube slightly longer than the calyx. G. Calyx villous-hirsute, somewhat glandular 47a. V. Gooddingii var. nepetifolia G. Calyx short-strigillose, not glandular 38. V. tampensis F. Corolla-tube at least one-half longer than the calyx. H. Plants prostrate-decumbent; leaves varying from cuneate to orbicular-ovate; calyx-teeth short (1-1.5 mm. long) 37. V. maritima H. Plants ascending, decumbent only at base; leaves ovate to elongate-ovate; calyx-teeth long (2-3 mm.) . . . .35. V. canadensis E. Leaves 3-cleft, incised-pinnatifid or bipinnatifid. I. Corolla-tube protruding well beyond the calyx. J. Corolla-tube approximately twice as long as the calyx. K. Plants somewhat hirsute; both surfaces of the leaves simi- lar in color; spikes elongating in age; calyx-teeth 2-3 mm. long 35. V. canadensis K. Plants hispid-hirsute; lower surface of leaves lighter than the upper; spikes compact at maturity; calyx- teeth rarely surpassing 2 mm. in length. L. Plants prostrate-decumbent; spikes few-flowered..S6. V. elegans L. Plants ascending, decumbent only at base; spikes many-flowered 36a. V. elegans var. asperata J. Corolla-tube one-third to one-half longer than the calyx. M. Floral bracts equalling or exceeding the calyx; calyx not glandular, hispid-hirsute. N. Leaves bipinnatifid, ultimate segments linear-oblong.. 40. V. bipinnatifida N. Leaves 3-cleft, with segments remotely incised or lobed, ultimate segments much broader than in the above. 40a. V. bipinnatifida var. latilobata M. Floral bracts shorter than the calyx; calyx for the most part somewhat glandular, hispid-hirsute to villous- pubescent. 312 ANNALS OF THE MISSOURI HOTANICAL GARDEN [Vol. 20 O. Leaves sessile, strigose-hispid 45. V. Andrieuxii 0. Leaves subsessile to short-petiolate, hispidulous-hirsute to hirtellous. P. Plants coarse; leaves usually 2.5-4 cm. long, hispid- ulous-hirsute; corolla-limb 6-12 mm. broad; nutlets 2.5-3 mm. long. Q. Calyx-teeth long (2-3 mm.). R. Plants usually tall (2-4 dm.); leaves bipinnatifid with ultimate segments lanceolate; calyx-teetl not especially conspicuous in mature fruit. i S. Calyx glandular J+l. V. ambrosifolia S. Calyx not glandular. ^/a. V. a?nl>rosifoliaf. cglarululosa II. Plants usually low (1-2 dm.); leaves trifid with segments more or less incised; calyx-teeth conspicuously long in mature fruit 43a. V. ciliata var. longidentuta Q. Calyx-teeth short (usually less than 2 mm.). T. Plants decumbent-ascending or prostrate, densely hirsute; calyx somewhat glandular, hispidulous-hirsute, scarcely viscid. U. Plants with decumbent-ascending loose habit; leaf-margin slightly revolute 43. V. ciliata V. Plants with prostrate compact habit; leaf- margin strongly revolute. .43b. V. ciliata var. pub era T. Plants ascending-erect, more or less hirsute; calyx densely glandular-hirsute and somewhat viscid-pubescent 42. V. Wrightii P. Plants slender; leaves smaller, 1-2.5 cm. long, hirtellous; corolla-limb 5 mm. broad; nutlets 2 mm. long 44, V. racemosa I. Corolla-tube slightly longer than the calyx. V. Calyx-teeth acute-subulate, short (less than half as long as the calyx-tube). W. Plants repent, slender; spikes few-flowered, scarcely pro- truding beyond the subtending leaves. X. Corolla-limb 5 7 mm. broad \6. V. tcucm folia X. Corolla-limb 3 mm. broad. .^tfa V. teucriifolia var. corollulata W. Plants decumbent-ascending, stouter; spikes many-flow- ered, Bubsessile or short-pedunculate. Y. Plants more or less hirsute, decumbent; corolla incon- spicuous, limb 3-5 mm. broad 48. V. pumila Y. Plants densely pilose-villous, ascending; corolla con- spicuous, limb 8-9 mm. broad 47. V. Gooddingii V. Calyx-teet h subulate-setaceous, elongate (approximately half as long as the calyx-tube). Z. Plants decumbent-ascending, villous-pubeseent; leaves ovate, trifid with the segments coarsely dentate. 49. V. setacca Z. Plants erect, sparsely hirsute; leaves narrow and elongate, bipinnatifid, with the segments linear 60. V. lilacina A. Leaves subauriculate and semiamplexicaul at the base 51. V. amoena 1933] PERRY — NORTH AMERICAN SPECIES OF VERBENA 313 33. V. quadrangulata Heller, Contr. Herb. Franklin & Marshall College (Bot. Expl. S. Texas) 1 : 84, pi. 6. 1895. V. pumila f. albiflora Standi. Field Mus. Publ. Bot. 4: 256. 1929. Helleranthus quadrangulatus Small, Fl. Southeast. U. S. ed. 1, 1011. 1903, anded. 2, 1913. Low plant more or less diffusely branched from the base; stems prostrate-ascending, rooting at the lower nodes, hirsute; leaves 1-3 cm, long, broadly ovate, with cuneate base contracted into a very short margined petiole, incised-pinnatifid or 3-cleft, with lobes incised, strigose-hirsute on both surfaces; spikes compact, sessile or nearly so, terminal; bracts somewhat shorter than the calyx, narrowly lanceolate, hirsute, ciliate; fruiting calyx 5 (-6) mm. long, hirsute, particularly along the ribs, lobes short, subulate-tipped; corolla- tube slightly longer than the calyx, practically glabrous without; corolla-limb 2.5-3 mm. broad, lobes emarginate; anthers unappendaged ; ovary sur- mounted by subhemispheric-angulate stylopodium; schizocarp constricted along the lines of cleavage; nutlets 4 mm. long, crowned with a smooth obtusish beak, shallowly reticulate- scrobiculate above, longitudinally striate and somewhat abruptly enlarged at base. Distribution: Texas and northern Mexico. Specimens examined: Texas: sandy beach and open flats, Rio Bravo del Norte, Sckott (NY); Devils River, Val Verde Co., 15 May 1913, Orcutt (MBG); creek between Del Rio and Comstock, 22 April 1925, Small & Wherry 12010 (NY); "Brackett" (? Brackettville), 22 Aug. 1900, Trelease 101 (MBG); Spofford Junction, 22 March 1900, Canby 193 (US); Uvalde, 28 April 1928, E. /. Palmer 83592 (G, MBG, NY); San Antonio, April 1922, Schulz 767 (US); 24 km. south of San Antonio, 28 April 1921, Schulz 475 (US); woods near Colorado River, near Wharton, 12 April 1925, Small & Wherry 11824 (NY); railroad north of Moore Station, Frio Co., 5 April 1901, Eggert (MBG); sandy open ground, Pleasanton, 16 May 1916, E. J. Palmer 1747 (MBG); El Jardin, 10 March 1924, Runyon G2S (US); Eagle Pass, April 1883, Havard (US); Kennedy- Beeville, 15 March 1929, Tharp 55SS (US); Kennedy-Portland, 17 March 1929, Tharp5610 (US); Corpus Christi Bay, Dec. 1879, Palmer 1046 (G); Corpus Christi, 10 March 1894, Heller 1388 (G, MBG, NY), type collection; San Diego, 1885-6, Croft 78 (NY); Kingsville, 27 March 1920, High 75 (MBG); between Laredo and Bejar, Feb. 1828, Berlandicr 1485 = 225 (G); Laredo, Feb. 1891, Dodge 154 (US); sandy ground, Laredo, 6 April 1901, Eggert (G, MBG); Laredo, 21 March 1903, Reverchon 3902 (G, MBG, NY, P, US); Laredo, 1913, Orcutt 5542, 5730 (MBG); San Antonio, 1926, Pagel 2208 (F); Guadalupe, 168 km. southwest of San Antonio, 314 ANNALS OF THE MISSOURI HOTANICAL GARDEN [Vol. 20 Sept. 1879, Palmer 2039 (G); chaparral near Ilarlingen, 6 April 1925, Small & Whirry 11903 (NY); Brazos Santiago, 1SS9, Nealley (F, US). Mexico: Tamaumiwh: east of Matamoros, May 1836, Herlandier 3018 = 1518 (MBG). Verbena quadrangulata is an anomalous species often confused with V. pumila on account of the strong similarity in habit, but is perhnps more closely related to V. delticola, the only other known North American species with a tendency toward develop- ing a nutlet with a beak. The style is enlarged at the base into a persistent subhemispherical body; hence, when the nutlets split apart they appear as if elongated at the apex into a beak-like appendage. Although the anthers are not glandular and the flowers not showy, the species seems to belong to the section Glandular in. 34. V. delticola Small, 22 n. sp. Stems decumbent to ascending, branched, more or less hirsute; leaves ovate-deltoid, 3-7 cm. long, with truncate-cuneate base narrowed into a margined petiole, obtusish or acutish, coarsely serrate-den late, often trilobed, usually thin, sparsely appressed- hirsute on both surfaces, trichomes above often with minute bulbous bases; spikes peduncled, fascicle-like in anthesis, be- coming elongated in fruit; bracts linear-attenuate, shorter than or equalling the calyx; calyx 7-8 mm. long, sparsely glandular, hirsute, lobes slender, subulate, unequal; corolla-tube protruding well beyond the calyx, pubescent or glabrate without; corolla- limb probably 10 mm. broad, segments emarginate; nutlets 3 mm. long, subcylindrical, reticulate from the apex to the defi- nitely broadened base; commissural face practically reaching the tip of the nutlet, muricately scabrous. " V. delticola Small, spec, nov., herbacea verisimiliter perennis; caulibus decum- bentibus vol asccndentibus ramosis plus minusve hirsutis; foliis ovato-deltoideis basi cuneata in petiolum alatum attenuatis 3-7 cm. longis obtusiusculis vel acutis grosse serrato-dentatis saepe trilobis utrinque sparse adpresso-hirsutis; spicis pedunculatis; bracteis lineari-attenuatis calyce brevioribus vel subaequantilms; calyce 7-8 mm. longo sparse glanduloto hirsute; calycis dentibus subulatis inaequalibus; corollae tubo exserto extus puhoscente vel glabrato; corollae limbo circiter 10 mm. lato segmontis omarginatis; coccis subcylindricis 3 mm. longis suporne reticulatis. — Collected at Las Palmas Ranch, vicinity of lbownsville, Texas, 1-5 Aug. 1921 Ferris & Duncan 3161 (NY), type. 1933] PERRY — NORTH AMERICAN SPECIES OF VERBENA 315 Distribution: Texas and Mexico. Specimens examined: Texas: Edinburg, Hooker 5999 (US); Samfordyce, 1927, Molby 7224 (US); chaparral near Harlingen, 16 April 1925, Small & Wherry 11901 (NY); Brazos Santiago, 1889, Nealley 117, 118 (US); Reynoldsville, Cameron Co., 11 April 1905, Lewton 141 (US); Las Palmas Ranch, vicinity of Brownsville, 1-5 Aug. 1921, Ferris & Duncan 3161 (MBG, NY type). Mexico: Gregg Walnut (MBG); river gravel, near Monterey, 11 July 1888, Pringle 2228 (G); Sierra Madre, near Monterey, 25 Aug. 1903, Pringle 11843 (G, US); Loma del Obispodo, Monterey, 1 Feb. 1907, Safford 1221 (US); Monterey, Guadalupe, May 1911, Arsene 6129 (G, MBG, NY, US); Monterey, 10-11 March 1923, Tharp 1826 (US). Tamatjlipas: Victoria, 23 May 1898, Nelson 4424 (G, US); vicinity of Victoria, 1 Feb.-9 April 1907, Palmer 39 (F, G, MBG, NY, US); San Fernando to Jimeney, 26-27 Feb. 1902, Nelson 6628 (? 6028) (G, MBG, NY, US); vicinity of Tampico, 1-31 Jan. 1910, Palmer 90 (F, G, MBG, NY, US). San Luis Potosi: La Hoya, Liebmann 11314 (US). Puebla: near Metlaltoyuca, 31 Jan. 1918, Goldman 42 (US). Vera Cruz: Wartenburg, near Tantoyuca, April 1858, Ervendberg 236 (G); Colipa, March 1841, Liebmann 11818 (US). Superficially, this species bears a strong resemblance to V. canadensis, but usually the leaves are not so deeply incised. Its distinctive character is found in the nutlet. Commonly, in the section Glandularia, the schizocarp is shallowly lobed at the apex; hence, the style appears to be attached in a very definite de- pression and ordinarily the commissural face does not reach the tip of the nutlet. In V. delticola, however, the depression is indefinite, the commissural face practically reaches the tip, and the seoarate nutlets viewed from the lateral or ventral surface tendency toward developing a beak. In Safford small Mem. Torr. Bot. Club 1894. V. Aubletia Jacq. Hort. Vind. 2: 82, pi. 176. 1772; Linn, f Suppl. 86. 1781; Bot. Mag. pi. 308. 1795; Bot. Reg. pi. 294 1818. V. Oblaetia Retz. Svenska Vet. Akad. Nya Stockh. Handl. 34 143, pi. 5. 1773. V. Obletia Medic. Act. Acad. Theod.-Palat. 3 : 194, pi. 7. 1775 V. longiflora Lam. Tab. Encyc. 1: 57. 1791. (Vol. 20 316 ANNALS OF THE MISSOURI IJOTANICAL GAItDEN V. rubra Salisb., Prodr. 71. 1790. V. Lamberti Sims in Bot. Mag. pi. 2200. 1821. V. Avbhlia var. Drummondi Lindl. Bot. Reg. ?>Z. 7 ##5. 1837. F. Lamberti var. rosea Sweet, Brit. Fl. Gard. 1 1. 4: 303. 1838. V. Drummondii Hort. ex (J. Don in Loud, llort. Brit. Suppl. 2:080. 1839. V. Drummondii (Lindl.) Baxt. ex Small, Fl. Southeast. U. S. ed. 1, 1011. 1903, and ed. 2, 1913. V. canadensis var. Lamberti Thell. Fl. Advent, de Montpellier, 428. 1912. Buchnera canadensis L. Mant. 88. 17G7. Billardicra cxylanata Moench, Method. 309. 1794. AnoviimoH caroliniensis Walt. II Carol. 104. 1788. (ilandvliuia caroliniensis J. F. Gmel. Syst. Veg. 2 2 : 920. 1791. 0. Aubletia Nutt. Trans. Am. Phil. Soc. 5: 184. 1837. Stems decumbent to ascending, rooting at the lower nodes, more or less branched, spreading-hirsute or glabrate; leaves ovate to elongate-ovate, 3-9 cm. long, 1.5-4 cm. broad, with truncate or euneate base narrowed into a margined petiole, in- cised or incised-pinnatifid or3-cleft, appressed-hirsute or glabrate on both surfaces; spikes pedunculate, fascicle-like in anthesis, becoming elongated in fruit; bracts shorter than (or occasionally as long as) the calyx, linear-attenuate, hirsute, usually ciliate; calyx glandular-hirsute, in fruit 10-13 mm. long, lobes very slender, subulate-setaceous, unequal, the posterior lobe much shorter; corolla-tube about twice as long as the calyx, glabrous without or finely pubescent or glandular; corolla-limb 11-15 mm. broad, segments emarginate; anther-glands not minute; schizo- carp at maturity constricted along the lines of cleavage; nutlets 3 (-3.5) mm. long, subcylindrical with slightly broadened base, reticulaie-srrobiculate; commissural face muricate-scabrous. Distribution: North Carolina to Florida, and west to Colorado and Texas. Specimens examined: North Carolina: roadsides, Kittroll, 11 April 1889, Sturtevant (MBG). South Carolina: Abbeville District, Hexamer A Maier (G). Georgia: by roads and margins of fields through lower Georgia and middle Florida, March 1843, Rugel (MBG). Florida: Fort King, Aldcn (NY); St. Nicholas, Duval Co., July 1898, Lighthipe 599 (NY); Mous Creek, St. John's Co., 15 April 1S79, /. D. Smith (US); rich woods in Daytona, 30 March 1906, Deam 1831 (US); Ocala, 4 April 1879, J. D. Smith (US). 1933] PERRY — NORTH AMERICAN SPECIES OF VERBENA 317 Alabama: above Tuscaloosa, 8 May 1875, E. A. Smith (US); above Tuscaloosa, 11 April 1892, Ward (US). Louisiana: without locality, Hale (G, US), Short (NY); Minden, 15 April 1901, Trelease (MBG); Winnfield, 13 April 1912, Petersen (NY); Natchitoches, 16 March 1915, E. J. Palmer 7001 (MBG); Chopin, Natchitoches Parish, 23 March 1915, E. J. Palmer 7071 (MBG); vicinity of Covington, 20 March 1920, Arsene 11982 (US); Jacksonville, Drummond (G); Cameron, 4 July 1903, Tracy 8707 (G, MBG, NY, US). Kentucky: Shelbyville, Flint (G). Tennessee: Cedar Glades, north of Lavergne, 4 May 1898, Eggert (MBG). Iowa: Council Bluffs, Vasey (G). Missouri: St. Louis, May 1833, Engelmann 884 (MBG); St. Louis, 8 July 1910, SherffSSS (G); Windsor Springs, 30 May 1890, Hitchcock (MBG); St. Louis Co., 16 May 1879, Eggert (MBG, US); Allenton, 27 April 1887, Eggert (MBG); west of Kimmswick, 13 April 1918, Drushel 3659 (MBG); Kimmswick, 30 April 1905, Johnson (MBG); Hillsboro, 24 May 1885, F. Wislizenus 285 (MBG); Crystal City, 19 Aug. 1 886, Eggert (MBG); sandy ground north of Crystal City, 6 May 1891, Eggert (MBG); Silica, 18 April 1896, Eggert (MBG); Shot Hill, near Selma, 30 May 1923, Greenman 4231 (MBG); Victoria, 10 May 1890, Hitchcock (MBG); Old Mines, Washington Co., 19 Aug. 192S, Kellogg 1958 (MBG); Blackwell, St. Francois Co., 18 April 1897, Trelease 714 (MBG); Central, 28 Aug. 1898, Trelease 1161 (MBG); banks of Pilot Knob Creek, 9 Sept. 1859, Engelmann (MBG); Pilot Knob, July 1867, Engelmann (MBG); Granite Mountain, Iron Co., 24 May 1918, Greenman 4076 (MBG); Ironton, 4 May 1923, E-pling 6134 (MBG); Shepard Mountain, Iron Co., 26 May 1918, Greenman 3870 (MBG); Shut In, Stout Creek, Iron Co., 28 May 1916, Drushel 2762 (MBG); near Silvermine, Madison Co., 20 May 1927, Greenman (MBG); Mine La Motte, 19 May 1927, Greenman (MBG); Grandin, 5 May 1901, Bush 818 (MBG); dry rocky banks, Shannon Co., 13 April 1889, Bush 1168 (MBG); Jerome, 5 April 1914, Kellogg (MBG); Jackson Co., 28 March 1864, Broadhead (MBG); Jackson Co., 28 May 1893, Bush 288 (G, MBG); Independence, 1 June 1895, Tindall (MBG); Dodson, 27 April 1904, Bush 1933 (MBG) ; rocky woods, Vale, 13 April 1908, Bush 4922 (MBG, US) ; barrens, Greenwood, 28 Oct. 1915, Bush 2901 (G, MBG, NY); Greenwood, 25 April 1911, Bush 6439 (MBG); Jasper Co., 16 Aug. 1893, Bush (MBG); Oronogo, 10 July 1910, E. J. Palmer 2996 (MBG); open banks, Webb City, 19 April 1903, E. J. Palmer 559 (MBG); Webb City, 4 May 1902, E. J. Palmer 804 (MBG); prairie, vicinity of Pearl, 22 Aug. 1912, Standley 9171 (US) ; Willard, 20 July 1919, Blankinship (P) ; Springfield, 31 July 1892, Dewart 74 (MBG); vicinity of Springfield, 29 Aug. 1911, Standley 8367 (US); barrens, Swan, 25 Sept. 1899, Bush 569 (MBG); Swan, 7-9 Oct. 1915, Eggles- ton 12240 (NY, US); Galena, Stone Co., 27 May 1914, E. J. Palmer 5682 (MBG); limestone slopes, "Bald Joe," Stone Co., 30 April 1924, E. J. Palmer 24616 (MBG); near Seligman, Barry Co., 4 April 1926, E. J. Palmer 29804 (MBG). Arkansas: Eureka Springs, 19 April 1899, Trelease (MBG); top of dolomite hill along White River, near Beaver, Carroll Co., 23 Oct. 1925, E. J. Palmer 29333 (G, MBG, NY); bluffs, Van Buren, 5 April 1929, Demaree 6411 (US); 14 km. west of Fort Smith, 1853-4, Bigelow (US); Fort Smith, 1 April 1928, Benke 4566 (G); Hot Springs, 31 Oct. 1899, Trelease (MBG); Malvern, Letterman (MBG, US); Prescott, 8 April 1900, Bush 545 (MBG); clay barrens, Fulton, Hempstead Co., 28 April 1914, E. J. Palmer 5407 (MBG); Red River, Pitcher (G). Kansas: Princeton, 19 May 1919, Street (P); between Olathe and Pleasanton, Miami Co., 18 June 1929, Rydberg & Imler 22 (MBG, NY); open woods, Cherokee [Vol. 20 318 ANNALS OF THE MISSOURI BOTANICAL GARDEN Co., 1896, Hitchcock 790 (G, MUG, NY, US); vicinity of Edna, 28 June 1929, Ryd- berg & 1 inter 379 (NY). Oklahoma: near Miami, Ottawa Co., 26 Aug. 1913, Stevens 2340 (G, MUG, NY, US); Sapulpa, 22 July 1K94, Bush 437 (G, MBG, NY, US); Perkins, 14 June 1893, Waugh 168 (MBG); upland woods, Muskogee, 30 June 1918, E. J. Palmer 14283 (MUG); Norman, 17 April 1915, Emig 416 (MUG, US); near Page, 20 May 1914, Blakeley 1471 (G); near Paul's Valley, Garvin Co., 19 April 1913, Stevens 109 (G, MUG); Colbert Station, 19 June 1891, Sheldon 20 (US); near Idabel, 18 May 1916, Houghton 8643 (G, MUG). Texas: Denison, 22 April 1904, Reverchon (MUG); Denison, 15 March 1904, lieverchon (MUG); wet places, T. & C. Junction, Bowie Co., 5 Sept. 1900, Eggert (G, MUG); sandy ground north of Longview, Gregg Co., 1 ( .) April 1899, Eggert (MBG); open ground, Longview, 21 April 1915, E. J. Palmer 7124 (MBG); sands, Dallas, Reverchon (G), 71,0 (MUG); Newland, near Dallas, 26 March 1901, Rever- chon S633 (MUG, NY); Grapeland, Houston Co., 28 May 1917, E. J. Palmer 1205C (MBG); Grapeland, 8 June 192(1, Thorp 888 (G, NY, US); Livingston, Polk Co., 10 April 191 1, E. ./. Palmer 5186 (MUG); Sabine River, 25 kin. north of Orange, 18 April 1899, Bray 66 (US); Dayton, Liberty Co., 21 May 1917, E. J. Palmer 11979 (MUG); sandy open ground, near Conroe, Montgomery Co., 15 April 1929, E. J. Palmer S3S19 (G, MUG, NY); Houston, Feb. 1842, Lindheimer (G); woods, Houston, 6 April 1872, Hall 435 (MUG, NY, P, US); Houston, Fisher (MUG), S3, 40, 47, 707 (US); Galveston, Van Huff (MUG). Colorado: Rocky Mountains, lat. 40-41', 1868, Vatey (G, MUG). Verbena canadensis is one of the most easily recognized of the North American species belonging to the section Glandularia. It has a large corolla with tube about twice as long as the calyx and, at maturity, an elongated spike with somewhat remote fruits and conspicuous calyx-teeth. The leaves are variously lobed or cleft. Although these variations have been used at sundry times to segregate species, they do not seem to be constant nor are they supported by definite geographic ranges, hence scarcely merit recognition. 36. V. elegans IIBK. Nov. (Jen. et Sp. 2: 273. 1818. V. mora n crisis Willd. ex Spreng. Syst. 2: 750. 1825. V. canadensis subsp. elegans Thell. Fl. Advent, de Montpellier, 428. I'M 2. l r . canadensis var. Ehrenbergii Thell. I. c. Stems prostrate-decumbent, tending to root at the older nodes, more or less branched, sparsely hispidulous-hirsute; leaves lanceolate-ovate with cuneate base narrowed into a margined petiole, 2-5 cm. long, incised-pinnatilid or trifid with the divisions less deeply cleft, somewhat appressed-hirsute on both surfaces, 1933] PERRY — NORTH AMERICAN SPECIES OF VERBENA 319 dark green above, lighter beneath, often with branches or fascicles of smaller leaves in the axils; spikes few-flowered, pedunculate, terminal, fascicle-like; bracts mostly one-half to two-thirds as long as the calyx, lanceolate, attenuate, glandular-hirsute, ciliate; calyx 8-10 mm. long, glandular-hirsute, teeth short, subulate and unequal; corolla-tube 15-18 mm. long, finely but sparsely pubescent without, throat gibbous; corolla-limb 10-12 mm. wide; nutlets subcylindrical, 3.5-4 mm. long, reticulate- scrobiculate chiefly on the upper half, lower part striate; com- missural face muriculate. Distribution: southern Mexico. Specimens examined: Mexico: Hidalgo: Sierra de Pachuca, 2700 m. alt., 22 July 1898, Pringle 6908 (ANSP, F, G, MBG, NY, P, US); Sierra de Pachuca, 21 and 22 July 1901, Rose & Hay 5556 (US); mountains, Pachuca, July 1905, Purpus 1433 (MBG, P); between Pachuca and Real del Monte, 19 July 1905, Rose, Painter & Rose 8668 (NY, US); Real del Monte, 9 May 1910, Clokey (MBG); Moran, Humboldt cfc Bonpland 4063 (Bot. Mus. Berl.-Dahl. type of V. moranensis, MBG phot.); under oaks and firs, Sierra de Ajusco, ca. 2800 m. alt., 26 Aug. 1902, Pringle 11092 (F, G, MBG, NY, US). Mexico: lomas de Santa Fe, Aug. 1927, Lyonnet 173 (US). Vera Cruz: near Chila, Distr. Ozuluama, April 1888, Seler 723 (G). Oaxaca: Tehuantepec, 1906, Gandoger (MBG). The above species is related to V. canadensis but is easily separable by its slender more or less prostrate habit, few-flowered compact spikes, and shorter calyx-teeth. The nutlets of the two are about the same size, but the reticulations in V. elegans are a little coarser than those in the more northern species. 36a* Var. asperata Perry, 23 n. var. Stems decumbent-ascending, hirsute-hispid; leaves variously cleft or lobed; spikes dense, many-flowered; calyx glandular, hispidulous-hirsute, teeth a little longer than in the species. Distribution: Sonora and Chihuahua to San Luis Potosi. Specimens examined: Mexico: Sonora: Sierra Madre, 15 Dec. 1890, Lumholtz 445 (G); Badehuachi, 2 Dec. 1890, Lumholtz 446 (G); Hermosillo, 1888, M. A. Crawford (G). 23 Var. asperata Perry, var. nov., caules ascendentes hirsuto-hispidi; foliis bi- pinnatifidis vel multifidis; spicis densis multifloris; calyce glanduloso hispidulo- hirsuto. — Collected at San Antonio, Coahuila, Mexico, 31 Aug. 1848, Gregg 355 (MBG), TYPE. 320 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 20 Chihuahua: south western Chihuahua, Aug. Nov. 1885, Palmer 295 (G, US). Duranoo: San Ramon, 21 Apr.-18 May 1900, Palmer 191 (G, MBG, NY, US). Coaihiii.a: Sierra de Paras, Oct. 1010, Purpus 4974 (F, G, MBG, US); 72 km. east of Saltillo, July 1880, Palmer 1052 (G, US); San Antonio, 31 Aug. 1848, Gregg 355 (MBG n Pi); Sierra Enearnaciore, 28 July 1S90, Nelson 3896 (US). San Luis Potom: near Morales, 187G, Sehaffner 716 (G); near San Luis Potosi, 1878, 1'nrnj & Palmer 720 (ANSP, F, MBG, US); Alvarez, 5-12 Sept. 1902, Palmer 61 (F, G, MBC, NY, US). This variety, as compared with the species, is stouter and more erect with coarser pubescence and larger spike. Palmer 191 differs from the oilier collections in having very shallowly lobed ovate leaves. Parry & Palmer 720 shows much variation in pubescence, some plants being scabrous, others scarcely so at all. Purpita 4 !> ?4 is lacking the coarse pubescence but appears more closely allied here than elsewhere. 37. V. maritima Small, Bull. N. Y. Bot. Card. 3: 436. 1905. Stems branched at base, decumbent or prostrate, sparingly pubescent or glabrate; leaves cuneate to orbicular-ovate, l-4(-6) cm. long, tapering into a margined petiole, incised-dentate or somewhat lobed, sparsely pubescent or glabrate on both surfaces; spikes terminal, pedunculate, fascicle-like in anthesis, becoming elongate in fruit; bracts linear-lanceolate, about one-half as long as the calyx, acuminate, pubescent, ciliate; fruiting calyx 10-13 mm. long, appressed-pubescent, often glandular, teeth short, slender, subulate; corolla-tube at least one-half longer than the calyx, pubescent without; corolla-limb 10-15 mm. broad; anthers with or without glands; nutlets subcylindric with broadened base, 4 mm, long, scrobiculate; commissure narrow, muricately scabrous. Distribution: Florida. Specimens examined: Florida: sandy ridges bordering the ocean, eastern Florida, Curtiss 1968 in part (G, MBC, NY); sandy soil, Merritt's Island, 9 Dec. 1929, Moldenke 219a (NY); near Cape Canaveral, 15 July 1896, Curtiss 5706 (G, MBG, NY, P, US); Cape Canaveral, 2-5 April 1904, Burgess 688 (NY); Fort Pierce, 8-9 April 1904, Burgess 713 (NY); sand dunes, Hobe Sound, 19 March 1921, Randolph 52 (5 ((J, US); Harksdale, 7 May 1918, E. J. Palmer I 151B (MBC TTPB). Nkw Mexico: collection of 1851-2, Wright t50S in part (G, NY). 24 V. tumidula IVrry, spec, nov., herbacea; caulibua decumbentibus vel ascen- dentibua ramosis piloso-hirsutis; foliis late ovatis basi cuneata in petioium alatum contractis trifidis segmentis in ei so- vel ^rosse crenato-dentatis subtus hirtellis supra adpresso-pubescentibua vel strigosis; spiois breviter pedunculatis paulo exsertis; bracteia ovato-lanceolatis acuminatis ciliatis oalyce fere paulo brevioribus; calyee 8-9 mm. longo basim inflato hirsuto el subli liter glanduloso; calycis dentibua 1.5 mm. longis; corollae tubo circiter 11 mm. longo extus puberulo; corollae limbo 8-10 mm. lato; connective) antherarum superiorum inappendiculato ; coccis subovoideis ."'» mm. longia dorso reticulato-scrobiculatis lateribus ventrico is et glabris; commissura subtiliter muriculata. — Collected at Barksdale, Texas, 7 May 1918, E. J. Palmer 18512 (MBG), TYPE. 1933] PEERY — NORTH AMERICAN SPECIES OF VERBENA 323 This species undoubtedly belongs to the section Glandularia, although apparently the anthers are glandless. Superficially, it resembles V. Gooddingii var. nepetifolia, but the spikes are not so large nor so showy. Its distinctive characters are the ventri- cose nutlets and the inflated fruiting calyx. The bracts, too, are broader and shorter than those of nearly related species of this group. 40. V. bipinnatifida Nutt. Jour. Acad. Nat. Sci. Phila. 2: 123. 1821; Schauer in DC. Prodr. 11 : 553. 1847. Glandularia bipinnatifida Nutt. Trans. Am. Phil. Soc. N.S. 5: 184. 1837. More or less diffusely branched from the base; stems loosely ascending, occasionally rooting at the lower nodes or from sub- terranean branches, hispid-hirsute; leaves petiolate, blades 2-6 cm. long, bipinnately parted or tripartite with divisions more or less bipinnatifid, lobes linear or oblong, appressed-hirsute on both surfaces, margin at times revolute; spikes pedunculate, fascicle-like in anthesis, becoming elongate in fruit; bracts mostly longer than the calyx, linear-subulate, hispid-hirsute, ciliate; fruiting calyx 8.5-10 mm. long, pubescent, hispid-hirsute along the nerves, lobes very slender, subulate-setaceous from a broader base, unequal; corolla- tube about one-half longer than the calyx, pubescent without; corolla-limb 8-10(-12) mm. broad, lobes emarginate; nutlets cylindric with slightly broadened base, mostly 3 mm. long, reticulate-scrobiculate above, striate toward the base; commissure muricately scabrous. Distribution: South Dakota to Alabama, westward to Arizona and northern Mexico. Specimens examined: Alabama: between Cahaba and Beloit, Dallas Co., 26 April 1927, Harper 15 (G, NY, US); on limestone outcrops along small creeks, near Demopolis, Marengo Co., 14 May 1925, E. J. Palmer 27209 (MBG); Spring Hill, 1918, Graves 536, 1946 (MBG). Mississippi: Ocean Springs, 1892, Skehan 47 (G). Louisiana: vicinity of Alexandria, 19 May 1899, Ball 401 (G, MBG, NY, US); Cotes Blanches, St. Mary Co., 17 July 1893, Langlois (MBG, US). Missouri: prairies, Upper Missouri, 21 June 1839, Geyer (US); Courtney, 6 June 1894, Bush 851 (G, MBG); Courtney, 13 June 1906, Bush 4029 (G, MBG, NY, US). Arkansas: "Red River/' Nuttall (G, NY), type collection; dry gravelly hills, Fulton, Hempstead Co., 17 June 1915, E. J. Palmer 8045 (MBG); near Homan, 10 324 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 20 June 1898, Eggert (MBG); along railroads, Miller Co., 19 June 1908, Eggcrt (NY); Texarkana, Aug. 1880, Lette.rman (MBG, NY). South DAKOTA! Fort Pierre, Nicollet's Northwest Expedition (US); Pierre, 2 Sept. 1891, T. A. Williams (MBG, NY); near McClure, June 1910, 0. E. White (MBG); Kennebee, Lyman Co., 1 July 1914, Over 8177 (US); Pine Ridge Reservation, Washabaugh Co., 22 Aug. 1911, Visher USB (F, NY); White River, 1G July 1896, T. A. Williams (NY); White River flood plain, 4 Aug. 1914, Over 2103 (US); Whilr River Valley, Shannon Co., 8 July 1911, Visher 2177 (NY); Fall River Falls, 18 June 1S92, Rydberg 985 (G, NY, US). Nebraska: Spencer, Boyd Co., 25 July 1893, Clements 2776 (G, US); Callaway, 4 July 19(11, Hates (G). Kansah: Cloud Co., 19 May 1SSS, Carlton (MBG); near Osborne City, 19 May 1894, Shear 88 (G, NY, US); Manhattan, 24 May 1901, A. Nelson 8245 (G, MBG, NY, US); plains, Ellis Co., 16 July 1895, Hilcheock S9S (G, MBG, NY, US); Smoky Hill, June 1867, Parry 157 (G, MBG); vicinity of Hays, 20 July 1929, Rydberg & Imler 1212 (NY), 1253 (MBG, NY); prairies, Medicine Lodge, 12 Sept. 1890, Smyth 306 (US); Morton Co., July 1891, Carlekm 177 (US); vicinity of Richfield, 20 Sept. 1912, Rose & Fitch 17105 (US). Oklahoma: Vinita, 3 Aug. 1877, Gurncy & Monell (MBG); Grant Co., 20 June 1899, While 11)7 (MBG); Woods Co., 29 June ?1899, White 165 (MBG); near Alva, Woods Co., 28 Sept. 1913, Stevens 2850 (G); near Whitehorse, 31 May 1913, Steven* 699 (G, MBG, NY, US); about 17 km. north of Boise City, Cimarron Co., 22 Aug. 1927, Stratton 449 (MBG); near Shattuck, Ellis Co., 20 May 1914, Clifton S085 (G, P); Hollis, Harmon Co., 21 June 1913, Stevens 1108 (G); 10 km. southwest of Hollis, 6 Aug. 1927, Stratton 817 (MBG); vicinity of Fort Sill, 27 April 1916, Clemens 11751 (MBG); on false Washita between Fort Cobb and Fort Arbuckle, 1868, Palmer 47 (NY); Arbuckle Mountains, Davis, 29 Oct. 191 1, Emig 895 (MBG); near Davis, 23 June 1917, Emig 788 (MBG); Mannsville, Johnston Co., April 1916, Griffith 8466 (G); Price's Fills, Murray Co., 30 April 1926, Stratton 10 (MBG); near Crusher Spur, Murray Co., 11 April 1913, Stevens 14 (O, MBG, NY, US); near Camp, Texas Co., 12 May 1913, Stevens 427 (G); Caddo, 22 June 1891, Sheldon (MBG), 48 (US). Texas: without data, Undheimer 282 (MBG), 289 (F, G, MBG, NY, US), 807 (G); Canadian, Hemphill Co., 10 Aug. 1900, Eggert (MBG); Canadian, July 1903, Howell lit {UX); vicinity of Terrell, Kaufman Co., 4 May 1904, Tyler (US); Dallas, Revcrchon 739 (MBG, US), 196B (F, G, MBG, NY, US); Dallas, Reverchon (G, NY, US); Dallas, 22 June 1899, Eggvrl (MBG); dry ground, near Garland, 24 June 1S99, Eggert (MBG); Fort Worth, Ruth 107 (F, G, MBG, NY, US); Weatherfonl, 26 May 1902, Tracy 7999 (F, G, MBG, NY, US); Coreicana, Navarro Co., 1 June 1915, E. J. Palmer 7821 (MBG); near Granbury, Hood Co., 4 May 1900, Eggert (MBG); Granbury, 4 Sept. 1914, E. J. Palmer 8610 (MBG); near Abilene, 7 June 1900, Eggert (MBG); Abilene, 19 May 1902, Tracy 8000 (F, MBG); Killeen, 1 Oct. 1891, Ward (US); Walker Co., 6-12 May 1910, Dixon 661 (F, P); near Austin, Travis Co., 18 April 1929, E. J. Palmer 383S9 (G, MBG); Crab Aj.pl.! Creek, Gillespie Co., Jermy I I (MBG); about 25 km. west of San Felipe, March 1844, Lindheimer 146 (MBG); San Marcos, 11 June 1S97, Treleaee (MBG); Comanche Spring, New Braunfels, Lindheimer 1072, 1078 (F, G, MBG, NY, US); northwest of New Braun- fels, 14 Sept. 1913, Penncll 5444 (NY); limestone soil mixed with sand, near Bracken, Groth 80 (F, G, NY, US); Rock Springs, 17 April 1930, Jones 26228 in part (MBG); Del Rio, 18 April 1930, Jones 26229 in part (P); Fort Clark, Kinney Co., Mearns 1252, 1274, 1894 (US); Eagle Pass, Rio Bravo del Norte, Feb.-March 1S52, Schott 1933] PERRY — NORTH AMERICAN SPECIES OF VERBENA 325 (NY); San Antonio de Bexar, Feb. 1828, Berlandier 1449 = 189 (G); between Trinity River and Bexar, June 1828, Berlandier 429 = ?1 749 (G) ; San Antonio, 4 April 1901, Eggert (G); San Antonio, 1897, Wilkinson 9 (MBG); San Antonio, 15 Aug. 1906, Ball 909 (US); vicinity of San Antonio, April 1919, von Schrenk (MBG); San Antonio to Austin, 18 April 1925, Small & Wherry 11968 (NY); 16 km. west of San Antonio, 8 June 1931, Moore & Steyermark 3001 (MBG); along Trinity River, near Liberty, 11 April 1925, Small & Wherry 11781 (NY); Columbia, 27 March 1900, Bush 449 (MBG, NY, US); Columbia, 3 May 1900, Bush 205 (MBG, US); Cuero, 22 March 1907, Howell 808 (US); prairies, near Victoria, 10 April 1900, Eggert (MBG); Victoria, 27 April 1905, Maxon 8808 (US); Victoria, 7 March 1916, E. J. Palmer 9098 (MBG); from Goliad to Bexar, May 1834, Berlandier 2428=998 (G, MBG). ?Montana: sandy soil along Yellowstone and upper Missouri, 1853-4, Hayden (MBG). New Mexico: without data, 1850-2, Wright 1502 (G, NY, US), 1504 in part (ANSP); below Dofiana, Parry, Bigelow, Wright, Schott (NY); Santa Fe, 20 July 1898, Earle 84 (MBG, NY); Roswell, Aug. 1900, Earle & Earle 526 (MBG, NY). Arizona: Bill Williams Mountain, 22 July 1898, MacDougal 817 (ANSP, F, G, NY); by streams of Santa Catalina Mountains, 13 April 1881, Pringle (G); Santa Rita Mountains, 31 July 1927, Peebles, Harrison & Kearney 4549 (US); Miller's Canyon, Huachuca Mountains, July 1909, Goodding 251, 495 (G, NY); road to Rustler's Park, Chiricahua Mountains, 18-19 June 1930, Goodman & Hitchcock 1211 (MBG); Barfoot Park, Chiricahua Mountains, 31 Aug. 1906, Blumer 1845 (F, G, MBG, NY). Mexico: Chihuahua: near Colonia Garcia, 21 July 1899, Townsend & Barber 189 (F, G, MBG, P, US); near Colonia Garcia, 1899, Nelson 6117 (G, US); Round Valley, Sierra Madre, 17 Sept. 1903, Jones (P). In the specimens from Arizona and Mexico, it should be noted that the pubescence is hispidulous and the spikes at maturity are, for the most part, dense rather than elongated. The typical form of the species is very easily recognized by the hispid-hirsute pubescence, particularly of the inflorescence, the lack of glands, the long floral bracts, the elongating spike, the deeply cut leaves, and a rather coarse habit. Apparently it intergrades with V. ciliata in northern Mexico, with V. ambrosi- folia in New Mexico, and at times with V. canadensis in the south- western part of its range. Nevertheless, in spite of the many intermediate phases, it seems preferable to maintain each group as specific entities. 40a. Var. latilobata Perry, 26 n. var. Leaves trifid with the segments remotely incised or lobed, 26 Var. latilobata Perry, var. nov., folia trifida; segmentis grosse incisis lobisve supra strigosis subtusque praesertim in nervis et venis hirtellis hirsutisve; spicis maturitate densis vel elongatis. — Collected between San Pedro and Fronteras, Sonora, Mexico, 20-24 SeDt. 1890. Hartman 906 (G). type. [Vol. 20 326 ANNALS OF THE MISSOXJRI BOTANICAL GARDEN strigillose above, hirtellous-hirsute, especially along the veins beneath; spike compact or somewhat elongated in age; bracts variable* in length, usually equalling or exceeding the calyx; floral and nutlet characters as in the species. Distribution: southern Arizona jind northwestern Mexico. Specimens examined: Akizona: Galiuro Mountains, 9 July 1894, Tourney (NY); Bisbee, Oct. 1908, Ooodding 87 (NY); Flagstaff, 17 June 1887, Meams (NY); Fort Iluachuca, Aug. 1893, Wilcox (NY); Fort Iluachuca, Sept. 1S94, Wilcox (US); Iluachuca Mountains, 29 June 5 July 1903, Griffiths 48. I (US); Iluachuca Mountains, 3 Sept. 1903, Jones (P); Miller's Canyon, Iluachuca Mountains, 26 Aug. 1910, Goodding 888 (G, NY); Santa Rita Mountains, 20 Sept.-4 Oct. 1902, Griffiths l (P, US); Trout (-reck, Charleston Mountains, about 1900 m. alt., 1926, Jaeger (P); Calientes, about 1400 in. alt,, 29 April 191 I, Jones (P); Kernan, Valley Meadow Wash, 28 April 1902, Goodding 645 (F, G, MHO, P), type collection. Aiuzona: without locality, 1891, MacDougal 60S (NY); Chloride, about 1350 m. alt., 15 April 1903, Jones (P); Ib.ckberry, 24 May 1884, J one* (F, P, US); Peach Springs, June 1884, Levimon (US); Peach Springs, April 1893, Wilson 99 (US); Kingman, 11 March 1912, Wooton (US); Skull Valley, about 1300 m. alt, 28 April 1903, J ones (P); Hillside, about 1100 m. alt., 1 May 1903, Jones (P); near Williams, 11 April IQOfi, Wilcox (US); Jerome Divide, 25 July 1921, W. W. Jones (MBG); east of Jerom<« Junction, 1 May 1908, Tidestrom 898 (US); Rio Verde, Fort Whipple, 30 Aug. IK65, Cooes & Palmer 4<>8 (MBG); Fort Verde, 1888, Mearns (NY); Natural Bridge, 23 April 1904, F. M. Chamberlain 56 (US); Bradshnw Mountains, 22 June 1892, Tourney 305a (US); 16 km. south of Black River, White Mountains, about 1800 m. all,, 22 June 1930, Goodman & Hitchcock 1297 (MB< J, NY); Galiuro Moun- tains, 29 July 1894, Tourney (US); Tucson, 16 May 1896, Zuck (NY, US); washes, near Baboquivari Mountains, 24 Feb. 1923, Hanson A1021 (F, MBG); Baboquivari Mountains, 10 March 1926, Thaeke.ru ); 716 (15); 757 (19); 755 Gandoger, M (40). GarlxT, A. P. (30). (38). Engclmann, ( ! — (10, 17, 19, 23, 31, 35, Garcia, P. I. 331 (40a). 41); 834 (35); 836 (17); 337 (10). Gardner, J. It. 583 (19). Engclmann, II. (19). Gare.schc, E. E. (19). Epling, C. C. 6446 (22); 6 184 (35). Ervendberg, L C. 153 (10); 236 (34). Gates, F. C. 9994 (19). Gates, F. C. & M. T. 10708 (16). Evans, VV. II (43b). Gattinger, A. (19). Everinaim, B. \V. .970 (19). Eyerdain, \V. .1. 201, 482 (14). Ezell, A. 6699 (8). Gershoy, A. 553 (0). Geyer, C. A. — (16, 40). Gibbes, L. R. — (2, 15). Fassett, N. C. & I lotchkiss, N. 8822 (10). Gilman, M. F. 1120 (26a). Fawcett, II. S. 12 (31). Faxon, G. E. Fendler, A. - Glatfelter, N. M. Gleason, H. A. (8, 15, 17). (19, 31); 377 (19); (16). (31); 686 (41); 557 (31); 594 (25); 697 in part (19, 20). Fernald, M. L. — (16, 31); 296 (16). Fernald, M. \,, Hunnewell, F. W. & Goodding, L. N. 37 (40a); 207 (31); Long, 11. 10262 (15). Fernald, M. L. & bong, B. 10268 (10). 7/5 (15); 1945 (16). Goldman, E. A. 42 (34); 292 (47a). Goldsmith, G. 119 (15). 251 (40); 834 (20a); 495 (40); 6\$5 (47); 553 (40a); 96$ (47a); 1022, 2812 (31). Fernald, M. L. & Parlin, J. E. 928 (15). Goodman, G. J. & Hitchcock, C. L. Fernald, M. L. & Pease, A. S. 25247 (10). Fernow, B. E. (47a). 1211 (40); 1297 (47); 1317 (31). Gotfredson, Mrs. A. 90 (16). Ferris, R. S. & Duncan, G. D. 2474 (41); Graham (11). 2607 (26b); 2647 (42); 2726 (25); Grant, M. L. 3078 (16). 3161 (34); 8268 (24); 8837 (8); '1/3 Graves, E. W. 5&5 (1); 536 (40); 00/ (32); (16). Fink, B. — Fisher, G. L. Oil); £51 (19). Witt (15); 1946 (40); 70/,7, f/W/, (19). G T (17). (II, 10, 35, 41, 40); Greene, 10. L. (23, 26, 42, 43b, 50); 19 (10); 83, 40, 47 (35); 122 (24); 168 45 (41); 77 (20); 860 (22). (5); 207 (43); 0/0 (24); 003 (43); Greenraan, J. M. — (35); 244, 245 (15); S86 (29a); 050 (25); 3/.V (43); 300 (12); 337 (46); 400, 005 (1); 707 (35). Flint, M. B. (35). Flynn, N. F. 69 (6). Forrer, A. (:<0, 40a). 898, 1273 (16); /S70 (15); 1877, 1879 (10); 7330 (15); 1382 (10); 7577 (15) 1968 (10); J03/, 3640 (17); 3766 (15) 3370, 4076 (35); 4125 (17); 403/ (35) 44/0 (15); 4678 (17). 1933) PERRY — NORTH AMERICAN SPECIES OF VERBENA 347 Greenman, J. M. & Greenman, M. T. Hartweg, T. 174 (28); 175 (7); 176 (43); 28 (25) ; 56 (47a) . 177 (29); 1924 (22). Gregg, J. — (8, 29, 43); 11 (7); 26, 57 Harvey, F. L. 61 (19); 62 (31); 1958 (19). (43); 95 (29); 202 (34); 243 (29); 257 Harwood, R. D. (42). (43); 265, 276 (7); 355 (36a); 406 Havard, V. — (25, 27, 31, 33, 42, 44, 48); (7); 483 (31); 545 (29); 628 (31); &?# 97 (42); /97 (27). (43); 686 (31); 34# (43); 7^0 (29); 752 Hayden, F. V. — (15, 16, 40); 10 (31). (34) ; 756 (43) ; 764 (5) ; 791 (29a) ; Haynes, D. (22). 823 (7); 1001 (11). Griffith, F. 8456 (40). Hedgcock, G. G. — (15, 16). Heller, A. A. — (15, 31); 638 (19); 1888 (33); 1419 (8); 1732 (29a); 5735, 5P20, 6778 (22); /#«* (16); /4503 (31). Heller, A. A. & E. G. 8586 (42). Henderson, L. F. 4070 (31). 1#00 (19); Griffiths, D. 1595 (25); 1919 (41a); *£M (47a); S4SJ (26a); 8489 (47a); 4244 (41); 4757 (43) 4^33 (40a); 5050 (19); 6190 (42); 55^0 (31). Griffiths & Hunter, /3 (31). Griffiths, D. & Thornber, J. J. 159 Herrick, C. L. 704 (42); 7/5 (26b). (40a). Grimes, E. M. 3716, 3738 (17); 3794 (15). Hertel, H. (15). W (16, 17, 35). Groth, B. H. A. 16 (29a); SO (40); 75 Heyde, E. T. 120, 477,580 (11); 610 (5). (8); 737 (24). Guidroz, H. 3 (15). Gurney, J. & Monell, J. T. Hale, T. J. Heyde, E. T. & Lux, E. 3018 (11); 3019 43 (40). Hicks (15). 245 (8) . (8, 15, 17, 19, 24, 35); High, M. M. 52 (8); 59 (43a); 76 (33). Hilgard, E. — (8, 32). - (31). Hall, E. — (19, 31); 393 (31); 428 (48); Hillman, F. H. - 429 (25); 480 (29a); 431 (48); 432 (8); Hitchcock, A. S. 433(1); 434 (24); 435 (35). Halsted, G. 175 (15). Hammond, E. W. 322 (22). Hansen, G. 964, 1823 (22); 2025 (5). (2, 14, 15, 16, 17, 32, 35); 269 (14); 393 (40); 790 (35); 791 (17); 972 (19); H29 (31). Hoffstetter, G. W Hanson, H. C. (48); A 146 (43b); Holm, T. (15). (15). (31). A147 (31); A148 (20); 322 (48); 651 Holzinger, I. M. — (19, 26, 41a). (42); 619 (25); 645 (26b). Hooker, L. H. 5999 (34). Hanson, H. C. & E. E. A1021 (47); Hough, W. 109 (43). A11S0 (26a). Harger, E. B. — (16, 17). Harper, R. M. 15 (40); 242 (14); 385 (17). Harris, J. A. C 16375 (47a). Harris, W. 9182 (2); 9937, JJS03 (14); 11969 (1). Harrison, G. J. 4778 (26a); 4397 (14). Houghton, H. W. 2570 (48); 3S22 (17); 3643 (35); 3346 (17); 4000 (19); 40/5 (15). House, H. D. 875, 1363 (17); 3276 (32). Howell, A. H. 64 (19); ///, 303 (40); 313, 862 (8). /£49 Hoyeradt, L. H. (17). Harrison, G. J. & Kearney, T. H. 8646 Humboldt, A. & Bonpland, A. (ID; (47a); 6796 (26a); 6144 (H); 6689 (26a). Harshberger, J. W. — (1, 2); 41 (46). 4063 (36); 4056 (12). Hyams, M. E. Jaeger, E. C. - (32). (47). Hartman, C. V. 94 (11); 60S (26a); 880, Jermy, G. — (48); 84, 86 (48); 182 (29a); 906 (40a). JS3 (40); 184 (48); 203 (8); 209 (48). 34S IVol. 20 ANNALS OF THE MISSOURI BOTANICAL GARDEN Jesup, H. (!. (16). Johnson, A. CJ. — (15, 16, 19, 35). Johnston, E. L. 507 (16); 986 (41). Johnston, I. M. — (14); 1820, 1608 (22). Lay bourns, Jones, M. K. — (7, 8, 11, 16, 19, 20, 22, Lea, M. C. Lansing, 0. E. — (16); 2805 (16); 2806 (15); 2810 (19); 2976, 3135 (17); 3164 (19); 8841, 3340 (16). W (6) (15). 23, 25, 26a, 40, 40a, 42, 43b, 47, 47a); Lee, D. W. 109 (4S). 88 (47a); 122 (41); 872 (26); 467 (42); Lehinann, F. C. 1608 (46). 487, 528 (16); 55^ (29); flty (41); Lsibsrg, J. B. 902 (31); 26'30 (16). f/.W (43); /;,S7 (16); 2/W3 (22); 8801 Lenmion, J. G. (43b); 8901 (47a); 5^73(31); 6026 (20); (28, 47, 47a); 2S57 (26a); 8075 (II). N,W.? (16); 23248 (4,X); ^3^4 (43); Leon, Frere — (6); 685 (14); 3910 (1). rOW (II); 25#W (43b); 26220 (IS); Leon, Frere & Fdinunds, Frere (6); •221 (25); 26222, 26223 (29a); 3/7.9 (14). 2tf2£$ in part (261)); 26225, 26226 Leonard, E. C. 3. «? (6). (29a); JL'/tf$7 (43b); 2 (31); 15277 (15). Letterman, G. W. — 35, 40). Lewton, F. L. 141 (34). Liebmann, F. M. 11308 (46); //3/S, Z/3/4" (34); 11316 (46); //»/« (13); 11828, 11329, 11330 (46); //&S5 (10). Lighthips, L. H. 699 (35). Lindheimer. F. (8, 14, 16, 24, 35); Kellogg, W. .38). Kelsey, J. A. 173 (16). Kempton & Collins (11). Kendall, M. L. Kennedy, (}. ( < (22). 15). Kennedy, P. B. 7032 (31); 70S3 (47a). Kerber, E. 255 (43); 3// (5). Killiam, (). L. 6988 (15). Killip, E. P. 8610 (5). King, 11. .17). Kirn, A. J. 2189 (14). Knause, < >. (15). Knowlton, C. 11. (16). KortholT, L (3). 145 (24); 146 (40); f&f (24); 166 (8); //AT (15); 232, 289 (40); 2#4 (29a); 3/// (40); 434 (48); 500 (29a); 601 (48); ff/3 (14); /07*, 1073 (40); /074 (29a); /075 (48); 1076 (8); /077 (14). Lloyd, C. E. 448 (11). Lloyd, F. E. 159 (43). Lloyd, F. E. & Tracy, S. M. — (6); 20 (3); 22 (14). Loomis, H. F. & Thackery, F. A. 910 (47a). Louis-Marie, Pere, 141 (16). Lumholtz, C. 445, 446 (36a). Lunsll, J. — (16, 31). Lyall, D. — (31). Lyon, 6 (14). Lyon net, E. — (7); 173 (36). Kreager, F. (). 469 (16); 474 (31); 475 Maobride, J. F. 200 (31); 30^ (16). ( 1 9) . Krig, J. T. 3187 (14). Kuntze, 0. 4 i (43); 2109 (5); 28444 (2H); 23788 (16); 233// (24). Macbride, J. F. & Payson, E. B. 781 (7); 850 (23). 4873 Lane, W. C. (15, 16). MacDougal, D. T. 73 (47a); 153 (7); 24.9 (2(0; 286 (31); 3/7 (40); 5/?/? (16); 505 (47). Lang, II. A. 102 (14). Langlois, A. B. — (1, 24, 40); 128 (24). MacFlwee, A. 76'0 (17); 873 (6); 882 (15). 1933] PERRY — NORTH AMERICAN SPECIES OF VERBENA 349 Mackenzie, K. K. 96 (8); 261 (19); 42/3 (17); 4746 (15). MacMillan, C. — Macoun, J. 24268 (31). McAtee, W. L. 1953 (8); 2053 (19). (19). (16, 31); 1305 (17); McClatchie, A. J. (14). McFarland, F. T. 92 (15); 109 (17). 2939a (6); 2946 (2); 2947 (14); 2984 (1); 4316 (16). Moore, J. A. & Steyermark, J. A. 3001 (40); 3005 (29a); 3009 (41); 3277 (26b); 3299 (25); SS#4 (42); 33H (27); 3622 (41). Morales, R. 786 (11). Morong, T. — (16, 31). McFarland, F. T. & Anderson, W. 253 Morris, E. L. — (15, 16). (19). Morton, Mrs. K. F. (48). McKelvey, S. D. 1710 (29a); 1756 (25); Moseley, E. L. — (17, 31). 2046 (26b). Manning, W. H. Muehlenberg, G. H. E. (17). (19, 31). Mueller, F. (11); 325 (43); 766, 887 Marble, D. W. 737 (2). Marsh, C. D. 14224 (42). Mathias, M. E. 545 (41). Mattern, E. S. & W. I. - - (38). Maxon, W. R. 3808 (40). Maxon, W. R. & Harvey, F. L. 7951 (5). Maxon, W. R., Harvey, F. L. & Valen- Munz, P. A. & Johnston, I. M. 11289 (14). (11); 1209 (43); 1215 (29); 1319 (43). Mulford, A. I. — (31); 37 (42); 39 (20). Munz, P. A. 616 (15); 617 (16); 1149 (26a); 1231 (41a); 1249 (43); 1470, 1607 (24); 2207, 6349, 6598 (31); 6734, 9295 (22). tine, 7356, 7478 (5). Maxon, W. R. & Killip, E. P. 569 (2). Munz, P. A., Johnston, I. M. & Harwood, R. D. 4252 (47). Mearns, E. A. (40a, 47); 108 (43); Murdoch, J. (40a) . 624 (31); 1013 (26a); 1116 (28); 1138 Nash, G. V. 601 (32); 1248 (14); 2470 (41a); 1237 (29a); 1252, 1274, 1394 (38). (40); 1456 (29a); JSS7 (26a); /304, Nealley, G. C. — (33); 117, 118 (34). /S>/4 (47a); 1918 (26a); 227S, ^S6 Nelson, A. 320 (31); 505, 535 (19); 1652 (41a); #027 (11); 2677 (41); 26,93, 5009 (31). Mell, C. D. (38). Mell, C. D. & Knopf, E. C. (31). Mellichamp, K. Meredith, H. B. (32). (6). Merrill, E. D. & Wilcox, E. N. 776 (31). Merton, E. C. 2042 (26a). Metcalfe, O. B. 126 (42); 137 (31); (31); 2258 (16); 7671 (31); 8245 (40); 8354 (20); 8489 (16). Nelson, E. W. 196, 223, 242 (46); 1021 (11); 1105 (12); 1513, 1895 (45); 1943 (7); 3896 (36a); 3920 (43); 4424 (34); 4559 (43); 4577 (7); 4593 (28); 4604 (43); 4856 (40a); 6H 7 (40); 0/0/ , 6271 (26a); 6023 (34); (5739 (41); 6858 (43). 6/2 (26); 897 (31); 955 (26); ZOOS (31); Nelson, E. W. & Goldman, E. A. 7123a 1090 (42); 1231 (41); 1568 (27). (47a); 7425 (21) Michener, C. A. & Bioletti, F. T. 123 Nelson, J. C. 1429 (22); 1804 (16); 2609 (23). Millspaugh, C. F. 865 (17). (22). Ness, H. (1, 3, 5, 7, 11, 17, 24, 31, Nicholas, (24). (11). Nichols, G. E. 163 (2). Mohr, C. — 32). Molby, E. 7224 (34). Moldenke, H. N. 148 (2); 212 (38); Nicollet's Northwest Expedition, Nicolas, (29). 219a, 549, 586 (37); 591, 599a, 972 (40). (14); 1039 (38); 1091 (32); 1184 (3); Nieuwland, J. A. 2685 (17). 1330 (15); 1339 (16). Moore, A. H. 2569 (16); 2374 (14); Norton, J. B. (6, 15, 17); 389 (15); 390 (16); 391 (19); 392 (31). [Vol. 20 350 ANNALS OF THE MISSOURI IIOTANICAL GARDEN Nuttall, T. Oakes, W. (17, 40). (6). Oersted, A. S. 11822, 11324 (5). Olney, S. T. (16). O'Neill, II. — (14); 98(1 (32); 6309 (37). Orcutt, C. H. (18, 31, 33, 47a); 909 (18); 1228 (31); 1801 (23); 1802 (14); 1803 (47a); 1546, 1*49 (31); 2736 (2); 3321 (13); 8488 (11); 3551 (46); 3950 (12); 4045 (11); 4146 (46a)j 51# (47a); /J^ (10); 5^f (33); 5/»55 (25); 5657 (43a); 57 J 7 (25); 57.W (33); 5792 (43a); 5S67 (8); 6096 (41); Wll (29a); 6772 (43b); 6184 (44); 6757 (41); 6235 (25). Ortega, J. G. 4% 15 (7); 454 1 (47a). - (15, 16, 20, 41); Osterhout, (1. E. — 8165 (40). Over, W. II. 2103 (40); 5/77 (40); 14S86 (19). Overholls, L (). — (15, 17). Pace, I,. £5 (8). Pagel, L. E. *j80S (33). 6789 (17); ff/Off (31); C/07 (19); 6510 (40); 700/, 707/, 7/24 (35); 7360 (8); 7556 (24); 7821 (40); #000 (15); 8045, 9098 (40); 0/// (8); 9188 (29a); 027S, 0.W (48); 9485 (8); 0550 (32); /0002 (29a); 10003 (48); /O0S7 (8); 105(18 (16); JOG*?* (14); i/7/5 (8); //&$3 48); //070 (35); 12001, 12038 (1); /2056 (35); 12496a (31); /5/75, /.^5/ (48); 18282 (25); /3^<9S (29a); /.S5/5 (39); 18562 (48); /356S (29a); 18564 (8); /S7S0 (25); /3,s'6/ (31); 14283 (35); Z5262 (17); 24616 (35); £7200 (40); *74ftS (14); 28738 (16); «fl.W» (2); 29838 (35); *&ttS5 (29a); £0/?0^ (35); 80523 (25); 30672 (42); 80791 (26b); 3///7 (31); 3//5, 1 ? (41a); 81192 (41); 333/0 (35); 367 (48); 33.W0 (40); AWS7 (25); 33583 (42); 33502 (33); 33604 (8); 33605, 33646 (29a); 84065 (26b); 34240 (41); 3*733 (17); 35235 (32); 37627 (19). Palmer, W. ,16). Palmer, Edward — (17, 26a, 41, 47, 50); Pammel, L. H. — (17, 19); 78 (16); 85 25 (29); 85 (43); 39 (34); 47 (40); 61 (36a); 52 (26a); 74 (43); 73 (8); 79 (43); 32 (29a); 90 (34); /S5 (28); Pammel, L. H. & Blackwood, R. E. (19); 187 (22); 272, 1699 (17); 1806 (16). /4/ in part (5); 141 in part, 163, 191 8589 (31); 3688 (16). (coll. <.f 1S98) (7); 191 (coll. of 1906) Parish, S. H. (14, 47a); 2819 (22); (36a); 20// (28); 2(!8 (7); 2S/ (43); 205 (36a); 803 (29); 307 (47a); 303 5333, 7/40 (14); /0J855 (31); 11590 (22). (7); 300 (22); 310 (23); 3/2 (26a); 326 Parish, W. F. 700, 197 (47). (48); 330 (coll. of 1876) (47); 839 Pariah, 8. B. AW. F. 969 (22); IO4S (14); (coll. of 1896) (11); S30H (26a); 2/7/ (31); 11143 (14). 34/ ^ (23); 342 (22); 345 (43); 856 Parker, C. F. — (6, 17, 31). (7); 364 (11); 897 (14); 401 (43); Purlin, J. E. 456 (28); 377 (50); 0// (28); /040 Parry, C. C. (16). (47a); 161 (40). (14); /042 (7); 1043 (8); /(>;4 (29a); Parry, C. C, Kigelow, J. M., Wright, C. 1046 (33); 1047 (29); /043 (31); /050 A Schott, A. — (26b, 31, 40, 44). (41); 705/ (34); 1052 (36a); //53 (11); Parry, C. C, & Lemmon, J. G. 842 r >7 2014a, 2019 (5); 2087 (10); 2033 (25); 2089 (33); iff ;0 (25); 2055 (43); 20* 252//u (22); 9699 (31); 6433 (38). Palmer, E. J- 197 (16); 703 (15, 16); (22). 4 (11); zo Parry, C. C. * Palmer, E. 717 (7); 7/3, 719 in part (46a); 7/0 in part (43); 720 (36a); 722 (28). /S5 (19); 2M (17); 804 (35); 454 (31); Patterson, II. N. (19). 569 (35); 7035 (16); 1747 (33); 2980 Patzky, (28). (19); 203/ (17); 2996 (35); 33/0 (17); Pease, A. S. & Long, B. 22350 (16). 4229 (15); 4232 (17); 4340, 4*77 (48); Peck, M. E. 8702 (22). 5044 (8); 5/35, 5407, 57/32 (35); Peebles, R. H. 5353 (43); 6755 (47a). 1933] PERRY — NORTH AMERICAN SPECIES OF VERBENA 351 Peebles, R. H., Harrison, G. J. & Redfield, R. 4 (H)- Kearney, T. H. 3790 (26a); 4U8 (31); Rehn, J. A. & Viereck, H. L. (42). 45/ f 9 W. 848 W. Perkins, J. R. 1024 (1); 102 (2). Peterson, N. F. (35). Phelps, O. P. 799 (16); 800 (15). Pilsbry, H. A. (42). Piper, C. V. 6160 (22). Pitcher, (35). Renson, C. 175 (11); 291 (5). Reverchon, J. — (15, 19, 35, 40); 42 (8); 118 (25); 732 (8); 734 (15); 735 (19); 786 (31); 737 in part (25, 29a); 738 (48) ; 739 (40) ; 740 (35) ; 834 in part, 1953 (25); 1961 (29a); 1962 (40); 1963 (48); £//3 (31); 2//7 (48); 2118, 2582 (24) ; 2533 (35) ; S502 (33) ; 3903 (29a) ; SS04, 4314 (25). Reynolds, M. C. (32). Pittier, H. 414 (29); 1909 (11). W (31). Plank, E. N. (19). Plaskett, R. A. 142 (22). Plummer, F. G. Pollard, C. L. • (42). (38); 1191 (14); 1312 (17). W (16). Porter, T. C. (6). Price, S. F. — (15, 17). W. W (26a). Riddell, J. L. 1268 (8). Ridgway, R. 2431 (19); 2831 (15). Riehl, N. — (16); 9 (31); 135 (15); 196 (19); 456 (17). Robinson, B. L. — (19); 26 (32); 94 (2); //S (6); **7 (2); 184 (15); /5S, /93 (16); 559 (15); 572 (16). Rojos, 44 (11)- Rolfs, P. H. — (31); 191 (32). Pringle, C. G. — (11, 14, 25, 26a, 28, 40, Rose, J. N. 1677 (11); 1763 (7); 2435, 47a); 54 (28); 270 (26a); ///7 (43); ^&W (7); 1948 (10); 2223 (34); 2813 (11); &?£7 (46); 3^7 (41); 3551 (46a); 4^50 (46); 4769 (12); 47S4 (29); 4577 (5); 4892 (11); 57/5 (7); 6539 2769 (43); JJ00S (47a); 16155 (50). Rose, J. N. & Fitch, W. R. 17105 (40); 17106, 17508 (31); /7504, 17 S01 (41). Rose, J. N., Fitch, W. R. & Parkhurst, 17717 (20). (28); 6908 (36); 7590 (28); 759/ (43); Rose, J. N. & Hay, R. 5310 (12); 5488 8534 (7); 9135 (28); 3S/2 (11); 93/5 (43); 9529 in part (7, 11); 11091 (43) //092 (36); //09S (5); 11843 (34) Z3/57 (41); 13158 (43); /S/59 (28) Purpus, C. A. 187 (46a); 195 (49); 4^7 (29); /094 (26b); 1095 (43); /4S3 (36); 3406 (13); 3407 (43); 45#4 in part (31, 43); 4974 (36a); 5451 (10); 55/5 (29); 6061 (47); 6413 (1). Quaintance, A. L. Rafinesque, C. S. Randolph, L. F. & F. R. (14). (31). (28); 5556 (36); 6049 (46); 6212 (29). Rose & Hough, 4484 (46). Rose, J. N. & Painter, J. H. 6431 (43); 7741 (29); 7742 (43); 7799 (7). Rose, J. N., Painter, J. H. & Rose, J. S. 8382 (7); 8495 (28); 3496 (11); 8668 (36) ; 3753 (7) ; 8823 (46a) ; 397/ (29) ; 9077, 9151 (5); 9162 (29); 9593 (43). Rose, J. N. & Rose, J. S. //S93 (43). Rose, J. N., Standley, P. C. & Russell, P. G. 12451, 12934 (7); 12745 in part (48); 12840 (47a); 13130, 18422, (32); 52 I3449 (37); 474 (15); 587, 645 (6); 688 (1); Rothrock, J. F. 1004 (3); 1012 (14). Rovirosa, J. N. (47a). (5). W (15, 31, 32). W Read, A. D. — (14); 19 (20). Ruano, J. M. 332, 403 (5). Reade, J. M. — Redfield, J. H. 6430 (17). (1). (15, 16); 522 (19); Rugel, F. (15, 31, 32, 35); 121 (6); 127 (17); 156 (14); 305 (38). Ruger, M. — (17, 31). 352 |Vol. 20 ANNALS OF THIO MISSOURI MOTANICA1, GARDEN Hunyon, R 628 (33); 629 (29»); 630 (43a); 1S60 (16). Busby, II. II. — / 181 (7); 331! (31). 23 (36); 7fcf (13); f/74, /373 (43); 4194, 434' (13); 45(50 (15). (20, 42); f*4 (46); Sesse, M. & Mocino, J. M. ,W (51). Seymour, A. B. — , 48, 49 (17); 50 (8). Ruth, A. — (li); 107 (40); 108 (8); /OS Seymour, F. C. 511 (16); 60/ (15); 1324 (31); //0 (48); 495, fits (31); 75/, 740, 765 (17); ,\v.v, #33 (15); 1289 (25). (16). Seymour, A. B. & Earle, F. S. 118 (32). Rydberg, P. A. (16); 154 (17); -932 Sevmour, A. B. & \\ : i ite, (31). (19); 934 (31); 986 (40); /.#« (Ml); Simmon, W. C. 3638 (11). /5/."> (16); t716 (15). Rydberg, I'. A. & Carlton, E. C. 7043 (31). Rydberg, P A. & Garrett, A. O. .920/ (28). Shear, C. L. 38 (40); 191 (19); 202 (16); 8577 (20). Sheldon, E. P. — (31, 40); 26 (35); 38 (31); # (40); 40 (17); 8202, 8856 CM); 1 1 167 (16). Rydberg, 1*. A. & Imler, R. II. 22 (35); SherfT, E. E. — (19); 333 (35); 1649 (16); 120 (17); 879 (35); 4 ; (19); 432 (15); JOGS (31); 1212, 1253 (40). Rydberg, P. A. & Vreeland, F. R. — 5676 (-11). St. John, Mrs. O. — (20). 1797 (31). Shimek, P. (S). Short, C. W. — (17, 19, 35). Shull, (1. II. 168 (16). Singer, J. W. 178 (17); 308 (15). Safford, \V. E. 216 (17); /22J (34); 1398 Sintenis, P. 1074, 2010 (14). (43). SafTonl, VV. 10. & Mosier, 210 (37). Salas, (S. 82 (5). Salazar, F. — (5, 7, 10, 12, 28). Saddling, J. II. & Lei berg, J. B. 34<> (31). Skohan, J. (14, 32); 20 (27); 46 (8); 47 (40); 109 (8). Small, J. K (6, 9, 15, 16, 17, 19, 31, •2); 2100, 2124 (37); 4020 (14); 6707 (3); 8469 (38); S50S, , (37); 8686, 8732 (38). Sandberg, J. II., MacDougal, D. T. Small, J. K. & Carter, J. J. 1072 (14); k Heller, A. A. 264, 972 (31). S;mford, S. N. (19). 1077, 2994 (37). Small, J. K., Carter, J. J. & Small, G. K. 8811 (37). Sarvis, J. T. 122 (31). Savage, T. 1-1., Cameron, F. E. & Small, , I. K. & Heller, A. A. — (9); 432 Lenocker, J. E. (16). (17); 484 (6). Schaffner, J. G. 716 (30a); 717 (43); Small, J. K. & Small, G. K. 4887, 4341, 718 (5); 719 (29); 720 (28). Schiede, C. .1. VV. 88 (11). Schoenfeldt, L. 2915 (7). Schott, A. (33, 40). von Schrenk, II. 48). Schuchert, C. Schuette, J. II. (6, 15, 16, 31, 40, (17). - (15, 19, 31). Selmlz, 10. I). 255 (20); 475 (33); 611), 4 ',03,4520 (14); 5422 (37). Small, J. K. & Wherry, E. T. 11774 (*); 11781 (40); 11813 (1); 11824 (33); 11826 (29a); 11884 (43a); 11901 (34); ll.')03 (33); 11968 (40); 11969, 11980 (29a); H003 (8); 12010 (33). Small, ,J. K. & Wilson, P. 1961 (37). Smith, C. L. 221 in part (43); 222 (5); 224 (11); /4?4 (46); /737 (11). 701 (24); 7',}/ (16). 31, 41, 41a, 42, 43a, 47, 47a); 187 (27); W (20); 303 (41); 364 (42); ^O.v (31); 61fi (41); 0# (20); 0SS1 (31); 88 7 (41a); 2SS0 (31); 8849 (431.); S«5* (26b). Wooion, E. (). & Standley, P. C. (31). Worthen, ( i. C. Wright, C. Wol.hor, II. J. (16, 10). While, M. 166 (40); 174 (16); /''< (40); 200 (31). White, (). E. (40). Wiggins, I. W. & Gillespie, D. K. 3.r; 257 Alopecurus 285 ambrosifolia 32(5 forma eglandulosa 328 amoena 341 Andrieuxii 333 an (j iisl if olia 282 arizonica 300, 337 A uldctia 315 var. Drummondi 316 Aubletia var. Lambert] 343 1933] PERRY — NORTH AMERICAN SPECIES OF VERBENA 355 Page barbata 342 bipinnatifida 323 var. latilobata 325 biserrata 268 bonariensis 254 bonariensis var. littoralis 257 var. rigida 253 bracteata 304 bracteosa 304 var. albiflora 304 var. brevibraeteata 304 I >rasiliensis 255 brevibraeteata 338 canadensis 315 canadensis var. Ehrenbergii 318 subsp. elegans 318 var. Lamberti 316 canescens 301 var. Roemeriana 302 canescens 304 var. neomexicana 296 caracasana 257 carnea 309 Carolina 268 caroliniana 268, 309 forma or var. polystachya 268 forma or var. recta 271 carolinensis 309 ciliata 330 var. longidentata 331 var. pubera 332 confinis 304 cuneifolia 286 delicatula 342 delticola 314 diffusa 275 domingensis 262 Drummondii 316 Ehrenbergiana 267 elegans 318 var. asperata 319 elongata 254 erinoides 341 exilis 334 Gooddingii 335 var. nepetif olia 337 Page gracilis 300 grandiflora 341 grandiflora 342 Halei 265 Hanseni 257 hastata 278 hastata var. oblongifolia 267 var. paniculata 278 forma rosea 278 var. pinnatifida 278 hirsuta 268 imbricata 304 incarnata 342 inconspicua 338 integrifolia 342 Lamberti 316 var. rosea 316 lasiostachys 290 leucanthcrnifolia 265 lilacina 340 litoralis 257 litoralis var. brasiliensis 255 var. caracasana 257 littoralis (3 leptostachya 257 a pyenostachya 257 longiflora 315 longifolia 272 Lucaeana 293 MacDougalii 288 MacDougalii mut. rosella 288 macrodonta 289 maritima 320 Matthesii 342 menthaefolia 263 mollis 268, 286 moranensis 318 neomexicana 296 var. hirtella 298 var. xylopoda 297 Oblaetia 315 Obletia 315 officinalis 262 officinalis var. hirsuta 296 Orcuttiana 284 paniculata 278 356 [Vol. 20, H)33| ANNALS OF THE MISSOURI BOTANICAL (GARDEN I \* \ ge var. pinuatifida 278 paucifiora 268 paucifolia 268 paucifolia 342 perennis 299 innnatijida 278 plicata 294 polystachya 268 proatrata 290 pubera 332 pulchvlla 332 pumila 33S pumila forma albijlora 313 quadrangular is 254 quadrangulata 313 racemosa 332 recta 271 remote 300 repena 342 rigcns 286 rigida 252 riparia 267 robusta 292 Rocnicriana 302 rubra 316 nulls 30 1 rugosa 282 scabra 272 scabra 272 scabrella 343 Pago setacea 339 srtosa 263 simplex 282 sphaerocarpa 256 spuria 262 s. Stamen; (a) gland-like appendage. Fig. 7. Pistil; (a) sterile style-lobe, (b) stigmatic lobe. Fig. 8. Nutlet; (a) lateral, (b) commissural, (c) dorsal surfaces. V. macrodonla and V. Wrightii are members of the sections Verbenaca and (Uanrtu- laria respectively. Here are illustrated the contrasting characters of the inflorescence in respect to (1) the size of the flowers, (2) the staminal appendage, (3) the sterile and the stigmatic lobes in relation to each other and (4) the commissural surfaces of the nutlets. Fig. 9. Diagrammatic longitudinal section of the flower of V. canadensis, showing the anatropous ovule with axile place itation. Figs. 10 17. Diagrammatic transverse sections through the flower of V. cana- densis to illustrate briefly the development of the carpels. Figs. 12-17 are of the ovary .alone. I ig. 10. ( ross-section through the receptacle and the subtending bract after the central cylinder has broken up into (a) calyeine, (b) corolline, (c) staminal and (d) carpellary traces. Fig. 1 1. Showing the calyx and the corolla free from the ovary. In the latter, two breaks in the tissue have appeared. Probably these are the ontogenetic locules of a bicarpellarv ovary. Fig. 12. Showing the two locules united. Figs. 13 17. Progressive steps in the development of the four ovules and their respective locules from the bicarpellary ovary. This is accomplished by the pro- trusion of the parietal placentae toward the centre bringing about axile placentation. Again the placentae appear to turn outward completing the development of the locules. Figs. 18-19. Transverse sections of V. bracteata, demonstrating the subtrigonous form of the schizocarp in the section Verbenaca as contrasted with the subcylindric form of the schizocarp in the section Glandularta, cf. figs. 1(3 and 17. Ann. Mo. Hot. Gaud., Vol. 20, 1933 Plate 13 1 3 i__ I 4b 10 4 4 A • * 5 6 8 7 9 11 8 8b 12 13 14 15 16 17 18 19 KERRY-NORTH AMKRICAN SPECIES OK VERBENA [Vol. 20, 1033] 360 ANNALS OF THE MISSOURI HOTANICAL GARDEN Explanation of Plate PLATE 14 Verbena macrodonta Perry. From the type speeimen, Nelson & Goldman 7^25, in the Missouri Botanical Garden Herbarium. Ann. Mo. Bot. Card., Vol. 20, 1933 Plate 14 ■ ■ • t - PERRY— NORTH AMERICAN SPECIES OF VERBENA I Vol. 20, 1933] 362 ANNALS OF THE MISSOURI BOTANICAL GARDEN Explanation of Plate PLATE 15 Wrhcmi Betucm Perry. From the type specimen, Purpus 195 \ in the Herbarium of Pomona College. Ann. Mo. Hot. Gaud., Vol. 20, L933 Plate 15 Tl^M i • * ^ ••> ' "1 MKM' i ^P> • r; »i9 tc - • »"» C A ' > J/ ' '* PERRY— NORTH AMERICAN SPECIES OF VERBENA NUTRIENT SOLUTIONS FOR ORCHIDS F. LYLE WYND Washingt In a previous paper ('33) the author reported the result of growing seedlings of Cattleya Trianae Linden & Rchb. f., on three-salt solutions of various ratios, all having a total osmotic concentration of one atmosphere. It was found that the best growth was associated with solutions having low concentrations of phosphate ions. All other ions were varied between wide limits with but little apparent effect. There is ample evidence in the literature of plant nutrition to support the belief that any set of nutrient salts at a given total concentration will exhibit more or less definite ratios for optimum growth, particularly if external conditions be kept constant. Since the effectiveness of any salt ratio depends upon the total concen- tration, it is not possible to compare the effects of specific ions in solutions differing in the total amount of salt present. The addition of extraneous ions may modify the permeability of the protoplasm and disturb the effectiveness of a given salt ratio. The solutions studied in the present work differ from each other in many respects, and the biological value of specific ions is therefore modified or obscured so that no comparison on the basis of ionic composition may be made. Such a comparison is possible only with the triangular series of salt ratios at a given total osmotic concentration. In the previous paper, the nutrient solution of La Garde ('29) was cited as a very favorable medium for the germination and growth of orchids, but its effectiveness could not be attributed to the specific concentrations of the potassium or phosphate ions. The present study was carried out to test this further, and to ascertain which of several published nutrient solutions were most satisfactory for the germination of orchid seedlings. The technique was the same as that described in the previous paper. The seeds were Cattleya Trianae Linden & Rchb. f. and from the same pod as those used for the triangular studies. The flasks were inoculated June 10, 1932, and the measurements were taken January 21, 1933. The molecular compositions of the solutions used are indicated in table i; their composition in parts Ann. Mo. Bot. Gabd., Vol. 20, 1933 (363) 3G4 [Vol. 20 ANNALS OF THE MISSOURI BOTANICAL GARDEN w W / 5 on O « 7 H O w O o H / O o <> 2 H f OO r ('UN) ON'IiN (KIII^I OSH'HN' K)"N z 10*0 'Od 8 H>l 1051 () 7 III z ( e ON)' J <> ? IIZ'OS*lV •ONM O'HS'OS^) r, ( f Od) ,1J ; ) o o CO *o ^o c •I CO •o *C CO Oi o (N o CO 3 00 CO CO CO o O iO *C CO CO Cj O c5 * CO CO 8 cu ►3o 8 w Q o 00 »o CO Q CO IQ co l^ o q t3 cu .3 " * o3 03 QQ O oj o O co cu 03 S3 1933] WYND — NUTRIENT SOLUTIONS FOR ORCHIDS 365 million, in table n: and the growth data, in table All received 1 cc. of a M/200 suspension of ferric phosphate >repared as described by Livingston ('19). TABLE II COMPOSITION OF THE SOLUTIONS IN PARTS PER MILLION OF THE NUTRIENT COMPONENTS Ca Mg K PC 4 S0 4 N0 3 NH 4 N (Total) CI La Garde 3 GO 98 287 69S 390 388 283 308 658 Crone 213 49 387 153 474 623 140 Kimdson 169 25 112 137 462 525 137 225 Sachs 310 49 387 307 474 623 140 302 Pfeffer 135 20 186 140 78 545 123 Shive A 144 360 703 1710 1440 448 101 Schimper 291 42 260 235 168 1071 241 260 Shive B 208 480 563 1368 1930 646 145 Knop 169 25 134 175 98 525 118 58 Tottingham 399 347 700 1235 1390 1547 348 Hansteen- Cranner 202 60 129 314 1098 322 250 458 Zinzadze 49 17 62 47 112 51 15 23 58 TABLE III GROWTH DATA, BASED UPON THE AVERAGE OF 25 SKKDLINGS . 1 pH at end Solution Height in Diameter in % Total pH at of rw* ^-^ *k» v*^*'*» microns microns salt planting experiment La Garde 4243 1590 .400 4.9 3.8 Crone 3760 1213 .225 5.5 4.5 Knudson 3360 1145 .200 5.0 4.5 Sachs 3268 1300 .300 5.1 4.5 Pfeffer 2953 1135 .150 5.0 4.6 Shive A 2908 1250 .700 4.8 4.5 Schimper 2835 1223 .345 5.1 4.5 Shive B 2793 1213 .812 4.8 4.4 Knop 2360 1145 .162 5.0 4.5 Tottingham 1868 900 .820 4.9 4.5 Hansteen- Cranner No growth No growth .295 4.9 3.9 Zinzadze No growth No growth | .052 5.1 3.7 The fact that no growth occurred on the solution of Hansteen- Cranner and of Zinzadze was probably due to instability of the 36G | Vol. 20 ANNALS OK TH M MISSOURI HOTANICAL GARDEN pH of these solutions as shown by tlieir high aeidity at the end of the experiment. In this connection it is interesting to note that 4200 3600 2100 2800 1400 700- T T T g 5 g 8 o Fig. 1. The relation of growth to calcium content. Zinzadze, Pfoffer, Shive A, Knop, K nudson, ITansteen-Cranner, Shive R, Crone, Schimper, Sachs, La Garde, Tottingham. 4200 3600 i.t.t 2100 1400 700 8 g o I T Fig. 2, The relation of growth to magnesium content. Ziniadze, Pfeffer, Knop, Knudson, Schimper, Sachs, Crone, Hansteen-Oanner, La Garde, Tottingham, Shive A, Shive B. 1933] WYND — NUTKIENT SOLUTIONS FOR ORCHIDS 367 4200 5500 300- 2100 1400 700- 8 8 O s o o Fig. 3. The relation of growth to potassium. Zinzadze, Knudson, Hansteen- Cranner, Knop, Pfeffer, Schimper, La Garde, Sachs, Crone, Shive B, Tottingham, Shive A. 4200- 3500 2B00 2X00 . 1400. 700 1 o s o o 8 o I— § s o Fig. 4. The relation of growth to phosphate. Zinzadze, Knudson, Pfeffer, Crone, Knop, Schimper, Sachs, Hansteen-Cranner, La Garde, Tottingham, Shive B, Shive A. [Vol. 20 368 ANNALS OF THK MISSOUKI HOTANICAIj (JAHDEN 4300 3800- 8800 8100 1400 700 8 g i o Fig. 5. The relation of growth to sulphate. Pfeffer, Knop, Zinzadze, Schimper, La (iarde, Knudson, Crone, Sachs, Hansteen-Cranner, Tottingham, Shive A, Slave IJ. 4200 6b00 ft R00 2100 1400 700 - T $ 8 O Fig. G. The relation between growth and nitrate. Zinzadze, La Garde, Shive A, Knop, Knudson, Pffefer, Crone, Sachs, Shive B, Schimper, Tottingham. La Garde's solution also was strongly acid at the end of the experiment, probably because of the greater growth of the numerous seedlings upon it. The use of ammonium carbonate by 19331 WYND — NUTRIENT SOLUTIONS FOR ORCHIDS 369 La Garde was probably for its buffer action against this unde- sirable change of acidity with growth. The effectiveness of the concentration of carbonate used as a buffering agent was tested by preparing La Garde's solution with and without carbonate and then comparing the titration curves obtained with N/100 hydrochloric acid. In both cases, the titration curves were 4200 3500 2800 2100 1400 700 j. O o o I x A A o o o to o Ol o o 1 4 Ol CI o Fig. 7. Relation between growth and ammonia. Zinzadze, Knudson, La Garde, Hansteen-Cranner. identical The solutions were also analyzed for carbonate after autoclaving, but no positive test could be obtained. The auto- claving at 20 pounds pressure for 20 minutes at the initial pH necessary (4.25) undoubtedly destroyed the small amount of carbonate present. Examination of table in shows that the best growth occurred on the solution of La Garde, followed by that on Crone's and 370 [Vol.20 ANNALS OK THK M1SSOUJU HOTANI( 1 AL HARDEN Knudson's. While the seedlings on La Garde's solution were conspicuously larger than those on Knudson's solution, the Knud- son seedlings showed a definitely superior root development. The Crone seedlings were actually larger than those on Knudson's solution, but they did not appear to be so green. Of all the solutions tested, we would regard (hose of La Garde and Knudson to be the most satisfactory. 4200 3600 2fl00 2100 1400 700 ■ i i CJl O 8 ro O o « (A CJl o Fig. 8. Relation between growth and total nitrogen. Zinzadze, Shive A, Knop, PfefTer, Crone, Sachs, Shive B, Knudson, Schimper, Hansteen-Cranner, La Garde, Tottingham. Tlie graphs indicate the comparative growth in relation to the concentrations in parts per million of each ion. It is apparent that the quality of the solutions is not related to a specific amount of any one ion. The author regards the superiority of La Garde's solution to be due to the particular complex of nutritional factors, and not to the specific effect of any particular ion. The nature of its superiority might well be a favorable condition of permea- bility of the cells produced by chemical means not yet understood. 933 J WYND — NUTRIENT SOLUTIONS FOR ORCHIDS 371 4200 3500 . • • 2100 1400 700 3 o CO to en 3 o 8 § o Fig. 9. Relation of growth to chlorine. Zinzadze, Knop, Schimper, Sachs, Hansteen-Cranner, La Garde. 4900 5500 8900 2100 14O0 700 1 o 3 O CO i 3 O Fig. 10. Relation of growth to per cent total salt. Zinzadze, Pfeffer, Knop f Knudson, Crone, Hansteen-Cranner, Sachs, Schimper, La Garde, Shive A, Totting- ham, Shive B. In all figures, the vertical distance represents the height of the seedlings in microns, and the horizontal distance represents the [Vol. 20, 1933] 372 ANNALS OF THE MISSOURI BOTANICAL GARDEN concentrations in parts per million of the ions indicated. The vertical lines representing the growth on the various media are listed from left to right. Conclusions 1. Seeda of Cattleya Trianae Linden ota, and the C. del Angelo, small valley of El Copey 1790 m., and El Roble 2670 in., on the following day. On January IS they passed Alto de la Baraja 2933 m., Rancho de las Vueltas, C. de las Vueltas 3019 m., and Ojo de Agua 2760 m.; on January 19 the Paramo de las Vueltas and the C. de Buena Vista 3299 m., and La Muerte at 3130 in.; on the 20th, southward along the slopes of Buena Vista to Lagunilla 1857 m., the Alto del Palmital at 1211 m., reaching El General on the evening of January 21 where they stayed until January 28, 1891, and whence Biolley returned to San Jose\ Pittier and Tonduz continued to Buenos Aires, Terraba, and Boruca, whence Pittier returned by a somewhat different route to San Marcos and San Jos6, arriving on February 23, while Tonduz stayed in the field until March 8, 1891. The following season Tonduz left San Jose on October 18, 1892, more or less retracing the journey of the previous season to Buenos Aires, thence down the 11. Grande de Ten-aba or Diquis, stopping some time at Boruca, finally down the river to the coast and back by sea to Puntarenas, stopping at several points, arriving on April 17, 1893. The material from these two trips furnished the bulk of material upon which Miiller Argau (1891, 1893) based his account of the lichens. These collections are to be found in the Herbier Miiller Argau, 1933] DODGE — LICHENS OF COSTA RICA. I 389 which was purchased by the Herbier Boissier and is now at the University de Geneve. I have been told that a duplicate set exists in Brussels, but apparently none was returned to the Museo Nacional in San Jose. C. Werckle, an engineer who resided in Costa Rica for many years, collected much in the vicinity of Cartago. His specimens were determined by Lindau and may be seen in Berlin and in Werckle's herbarium at the Museo Nacional de Costa Rica. Werckle' was not at all discriminating in selecting samples for Lindau, so that most of the material in San Jose needed revision. The Calverts (1917) spent a year, 1909-1910, in Costa Rica and have given us a good account of its natural history. They brought back some plants which are in the herbarium of the University of Pennsylvania, but I have seen only one lichen from them (at the Farlow Herbarium). Maxon and assistants have visited Costa Rica for ferns and brought back some lichens which were sent to G. K. Merrill for study and are now to be found in the Farlow Herbarium (which acquired the Merrill herbarium) and in the National Herbarium of the Smithsonian Institution at Washington. Paul C. Standley, in 1924 and again in 1925-6, in company with Juvenal Valerio Rodriguez and Ruben Torres Rojas, made very extensive collections of lichens although his primary interest was angiosperms. These lichens were likewise sent to G. K. Merrill who was studying them at the time of his death. As curator of the Farlow Herbarium I continued this study in preparation for my trip to Costa Rica and am here reporting these collections. In 1925, during a two-weeks' visit, I secured a large amount of material from Cerro Carpintera and again in 1929-30 I secured several thousand specimens of cryptogams which are still being studied. The most complete set of these plants is in my personal herbarium and the next set, lacking only a few numbers of the commoner species, is at the Farlow Herbarium. Karl Danielson, an assistant to H. E. Stork, in 1928 collected a few numbers of lichens which are now at the University of Michigan. These are also reported here for the first time, duplicates being found in the Farlow Herbarium and in my own herbarium. 390 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 20 In 1920 and 1930 the Costa Rican government botanist, Alberto M. Brenes, collected many lichens in the vicinity of San Ramon and a few from other parts of the country. These were also turned over to me for study and are to be found in the Museo National, with duplicates in my own herbarium and in the Farlow Herbarium. The Oesterreicliische Costa Rica Expedition, under the leader- ship of Otto Porsch (1932) with G. Cufodontis as assistant botanist, spent the late verano and early invierno in Costa Rica, visiting much the same territory as I did. If lichens were collected they were probably studied by Zahlbruckner but I have seen no publication on this group, although many of the other groups are already published. Floristics It is too early to draw any very definite conclusions regarding the distribution of plants and the affinities of the flora, but certain typical distributions are evident. Families and genera which are abundant in the northern United States and are often widespread in the North Temperate Zone are usually found at the higher altitudes and reach their southern limits in the mountains of Colombia, except for oc- casional isolated species farther south. Such an example is the Peltigcraceae, which are widespread and abundant in the northern part of the United States. Only two genera, Nephroma and Peltujrra, reach the American tropics. Nephroma is rep- resented by a single species seen in only one restricted locality in Costa Rica. Peltigera is represented by about eight species, mostly strictly tropical with only one undescribed species reaching Magellanes, Chile. Another typical example is a northern family, which is char- acteristic of the Atlantic Coastal Plain, the Collemaceae. Here genera are mostly common to the eastern and western hemispheres but subgenera and species are much more restricted in distri- bution. These are mostly found on the coastal plains of Costa Rica, being characteristic on the plains of Guanacaste and occasional on the Atlantic Coastal Plain but not yet reported from Panama. The species belonging in these genera from the 1933] DODGE — LICHENS OF COSTA RICA. I 391 higher elevations have relations with those of the mountains of Colombia and not with those north of the Isthmus of Nicaragua. A third type of distribution is found in those families which are predominantly southern, as in the Stictaceae. Here we find them highly developed and characteristic in the highlands, relatively rare in the lowlands. The genera and subgenera are world-wide in distribution and the species also have wide ranges, becoming less frequent in numbers and in species in the North Temperate Zone. In this group, however, species of the moun- tains of Costa Rica may extend into North America along the Atlantic Coastal Plain, even as far as the Avalon peninsula of Newfoundland, while the species of the Coastal Plain in Costa Rica are southern in their ranges. Finally some cosmopolitan families in which it is difficult to determine a center of distribution have genera or subgenera which are strictly limited to tropical America. Further consideration of problems of distribution may be deferred until the systematic enumeration is complete. In general the affinities of the flora seem to be with eastern Brasil but since we have very few collections available from the inter- mediate mountain ranges of eastern Colombia and southern Venezuela and British Guiana, it is likely that stations of many of these species will eventually be discovered in these regions. Relatively few species are common to Mexico and Costa Rica or to Peru and Costa Rica, and practically no Costa Rican species extend as far south as Chile. In the following systematic enumeration I have included in the keys to species all those which have been described from tropical America, whether they have been found in Costa Rica or not. I have not included species which were not originally described from this area, although they may have been reported from it, since often these reports have been based on misdeter- mined specimens. While it is evident that political boundaries do not always coincide with natural floristic divisions, for the purposes of this study tropical America may be defined as Mexico and the West Indies southward to Peru, Bolivia, Para- guay, Brasil, and Uruguay. These are quite natural in the pres- ent state of our knowledge, as the lichen flora of northern [Vol. 20 392 ANNALS OF THE MISSOURI BOTANICAL GARDEN Mexico which might be expected to show affinities with that of the southwestern United States is practically unknown. Simi- larly, that of northern Argentina, which might be expected to be similar to that of adjacent Paraguay, has been little explored for cryptogams. Northern Chile, largely desert, has also been little studied, while the flora south of the desert has relatively few species which have been reported from tropical America as here defined. The greatest difficulty occurs in the exclusion of southern Florida where occur many species which are tropical rather than northern in their affinities. Fortunately for our purposes comparatively few species have been described from this region. KEY TO TROPICAL AMERICAN FAMILIES OK FOLIOSE AND EUUTICOSE LICHENS Apothecia more or less exposed, paraphyses growing above the asci to form a tissue, nsci early evanescent, leaving an enclosed dusty spore mass (maaedium) CONIOCARPINEAE Thallus foliose or fruticose, corticate, apothecia sessile Sphaerophouaceae Apothecia linear, elongate-elliptic or angular, not forming a mazedium. . . . GRAPI1IDINEAE Thallus fruticose, erect or dependent, with a basal layer attached to the substrate, corticate [not yet reported between Lower California and Prtu|; maritime Roccellaceae Apothecia circular, not forming a mazedium CYCLOCARPINEAE Thallus strictly crustose, but fruiting structures appearing fruticose owing to the proliferation of the apothecia from the margins, easily mistaken for a hepatic on cursory examination; Guiana Poly stroma Thallus loosely byssine from filaments of Cladophoru or Tn ntepohlia, apo- thecia with light-colored parathecium Coemhioniaceae Thallus filiform or dwarf fruticose, or squamose, very rarely small foliose [deferred for further treatment with the crustose species]. Algal symbiont Scytonema or Stigonema f apothecia more or less sunk in the thallus, small and easily overlooked Ephebaceae Algal s\ mbiont Chroococcaceae Pyuenopsidaceae Algal symbiont Ixivularia Lhhinaceae Thallus gelatinous, swelling greatly when moistened, dwarf fruticose, squamose or large foliose, algal symbiont usually Nostoc Collemaceae Thallus definitely foliose or fruticose, or if small, the form of the thallus not due to the algal symbiont. Hypothallus and rhizoids highly developed, thallus squamose to small foliose, upper surface corticate, algal symbiont usually Nostoc. . Pannakiaceae Hypothallus evanescent. Thallus large foliose, algal symbionts Nostocacene or Palmellaceae, spores fusiform to acicular, 2 or more celled. 1933] DODGE — LICHENS OF COSTA RICA. I 393 Both surfaces corticate, lower surface interrupted by more or less highly developed breathing pores, apothecia with well-de- veloped parathecia or amphithecia, stipitate or sessile Stictaceae Only upper cortex developed, the lower surface tomentose with more or less highly developed network of veins and tufted rhizoids, apothecia attached to the thallus over their whole under surface without true parathecium or amphithecium although surrounded by the torn remains of the tissue cover- ing the young apothecium Peltigeraceae Thallus squamose or small foliose, only upper surface corticate, algal symbiont Pleurococcns, apothecia with well-developed light- colored parathecium, spores various. No secondary thallus or podetia present, apothecia sessile Phyllopsoraceae Secondary thallus or podetia present, varying from simple stalk of apothecium without algae to highly developed branched or infundibuliform structures, primary thallus squamose or small foliose, often evanescent or degenerate, perhaps crus- tose in some species Cladoniaceae Thallus large foliose or fruticose, algal symbiont Pleurococcus, spores one- or two-celled or muriform, never elongate-fusiform to acicular. Apothecia with black parathecium, spores one-celled [in Gyrophora haplocarpa from Peru, the only species of the family so far re- * ported] Gyrophoraceae Apothecia with bright-colored amphithecia. Spores not placodiomorphous. Thallus foliose, spores one-celled Parmeliaceae Thallus fruticose Usneaceae Spores placodiomorphous. Spores hyaline, thallus usually bright yellow, at least the epithecium orange Theloschistace ae Spores brown, thallus usually more or less glaucous, epithe- cium brown or pruinose Physciaceae Sphaerophoraceae Thallus foliose or fruticose, corticate on both surfaces or the lower surface incompletely corticate, with Protococcus. Apo- thecia sessile on the margin, or on the lower side of the thallus, open at first or enclosed by an amphithecium. Only two genera have been reported from tropical America, both fruticose with solid axis and terminal apothecium. Aero- scyphus sphaerophoroides Lev. without an amphithecium has been reported from Mexico and Peru but has not been found in Costa Rica. Sphaerophorus is typically Subarctic-Antarctic in distribution, coming southward on the higher mountains with [Vol. 20 394 ANNALS OF THE MISSOURI UOTANICAL GARDEN a single species on the higher mountains of tropical America. The whole family is greatly in need of revision. Spiiahrophorus melanocarpus (Sw.) DC. apud Lam. & DC, Fl. Franc, ed. 2, 6: 178. 1805, excl. syn. Lichen vidanocarpusSwurtz, Nova (Jen. Sp. PI. Prodr. 147. 1788. Type: Jamaica, Swartz. Thallus ashy glaucescent or pale above, pale below, partly terete, partly compressed, smooth, not rugose, highly branched, 1-6 cm. high, primary branches 1-4 mm. thick, KOH yellow above, KOH - below, CaOCL 2 - ; cortex 20 \i thick, subpellucid, of thick-walled conglutinate irregularly woven hyphae; medulla I — , composed of hyphae 4-8 jjl thick; mazedium oblique in the ends of the thicker terete branches, becoming disciform, exciple lacerate, ascospores 7-11 y. in diameter. Spermatogonia immersed rface of the branches lening by a blackened verruciform mouth; spermatia oblong k 3 [i [description from Nylander and Vainio). The Costa Rican material does not altogether agree with th< above description, but there is so much number of species and their relationships. Cartago: P. V. Turrialba, 2000-2 400 m., Sta ndlcy 86818. San Josr: 1,. la Chonta, n. e. Sta. Maria de Dota, 2000 2100 m., Standley 42290; C. tie las Vueltas, 2700-3000 m., Standley & J. Valerio 4888i. Heredia: C. Zurquf, 2000 2400 m., Standi, y A J. Valerio 50436. ROCCELLACEAE This strictly maritime family has not yet been reported between Lower California and Peru, probably because no suitable mari- time habitats have been visited by botanists, for the group is abundant in such strictly tropical habitats as the Galapagos Islands. I did not have an opportunity to visit such myself, but had I been able to visit the cliffs along the various headlands between l'untarenas and Golfo Dulce, it is possible that I might have secured some representatives of this family. ( yOENOG* )NI ACEAE Thallus spongy byssoid, either adnate or forming dimidiate shelving masses, homoeomerous with Trentepohlia or Cladophora, 1933] DODGE — LICHENS OF COSTA RICA. I 395 whose filaments are partially surrounded by hyphae. Apothecia with pseudoparenchymatous parathecium ; asci 8-spored; spores hyaline, one- or two-celled; spermatia exobasidial. Thallus with Trentepohlia Coenogonium Thallus with Cladophora, apothecia unknown Racodium COENOGONIUM Ehrenb. Coenogonium Ehrenberg apud Nees ab Esenbeck, Horae Physicae Berol. 120. 1820. Thallus loosely spongy byssoid, either adnate or forming dimidiate shelving masses (suggesting a thin species of Poly- stictus), homoeomerous with Trentepohlia which is partially surrounded by a network of hyphae. Apothecia scattered on the upper surface, scutiform, usually with a short stipe, with a parathecium of thin-walled pseudoparenchyma without a me- dulla; paraphyses unbranched, often with swollen tips; asci 8- spored; spores hyaline, fusiform, ellipsoidal to elongate, one- or two-celled; spermagonia spherical, spermatia exobasidial, fusi- form, straight; sperm atiophores mixed with anaphyses. Clements segregated the species of Coenogonium with uni- cellular spores as Holocoenis. However, his proposal has not been followed by other authors, although Vainio and Zahlbruckner have proposed sections on this basis, Vainio using Coenobiatora and C oenobiatorina , while Zahlbruckner incorrectly adopted Holocoenis and C 'oenobiatorina . This family is very aberrant among lichens in several respects and perhaps would be better dropped and the species distributed among the fungi and algae. It seems more logical to regard the group as a case of parasitism of fungi on algae, as neither fungus nor alga seems to influence the development of the other, and there has been no further evolution resulting from the acquisition of a photosynthetic unit by the fungus. Taking this view, one would recognize about four or perhaps five species of fungi from tropical America, one species with unicellular spores about 6-10 x 2.5-4 [x belonging in Patinella; and three or four species with 2-celled spores belonging in Orbilia or a segregate from that genus. Among the algae one would recognize a dozen or more species of Trentepohlia which are often subject to the attacks [Vol. 20 396 ANNALS OF THE MISSOURI BOTANICAL GARDEN of Patinella and Orbilia. In the following key to the tropical American species of Coenogoniwn, as it is commonly understood by lichenologists, it will be noted that out of nineteen names proposed, four were based on specimens in which no apothecia were found. Four other species originally described in Coeno- gonium have already been transferred to Trentepohlia. I have carefully investigated the ascocarps of the fertile Costa Rican specimens and can find few characters which would separate species from each other, except spore size and septation. For lack of time to investigate this question thoroughly, I have here followed the traditional arrangement, as to do otherwise would involve serious nomenclatorial difficulties. To adopt the view that this group should be excluded from the lichens would invalidate Coenogonium, as based on a combination of characters of parasite and host and involve re 'description of its species in Patinella, Orbilia, and Trentepohlia. KEY TO THE TROPICAL AMERICAN SPECIES OF COENOGONIUM Apotheeial disc livid, cell walls brownish; spores unicellular; algae 11-16 m in diameter; French Guiana C. Leprieurii Apothecia earneous, yellowish or waxy white, never livid. Thallus dimidiate, not adnate to the substrate*. Thallus (hick, rigid, with white subsilky villum, zonate, sterile; Brasil. . C. Echinus Thallus thinner. Filaments 7-9 m in diam., cells not very distinct, greenish, sub- diiTuse, stratose C. subvirescens Filaments 12 \x in diam., rigid; Brasil C. Linkii Filaments thicker, 16-20 /i in diam., thallus less rigid. Cells 4-5 times as long as broad; Mexico C. confervoides Cells about twice as long as broad; spores 7 S.5 x 2 13 /i; Brasil. . . . C. arrocephalum Filaments and paraphyses thick, imperfectly known C. andinum Thallus adnate to effused. Filaments moniliform, 18 m in diam.; thallus glaucous green to tawny brown, margins paler; spores 2-celled, 3-4 times as long as broad; Cuba C. nionilifortne Filaments partially moniliform, partly cylindrical, cells 20-25 \x in diam.; Costa Rica C. lulerotrichum Filaments uniformly cylindrical. Filaments 23-36 m in diam.; striate apothecia 0.6-0.7 mm. in diam.; spores 11-15 x 25-35 /*, 2-celled C. disjunctum Filaments 20-28 n in diam.; spores unicellular, 10x3 n; Bor- bonia or Louisiana C. inter po.situm 1933] DODGE — LICHENS OF COSTA RICA. I 397 Filaments 17-20 m in diam.; cells 30-50 /i long; spores 8 x 2.5 ju, 2- celled; Costa Rica C. interponendam Filaments 15-17 m in diam.; thallus ashy green, pulvinate, up to 2 cm. broad; Peru C. pulvinatum Filaments 12-16 m in diam.; thallus yellowish; spores 8-10 x 3 /*; Colombia C. interplexum Filaments 12-14 m in diam.; thallus yellowish to green; St. Vincent C. Leprieurii v. panniforme Filaments 8-11 p in diam., conglutinate in fascicles 70-90 m in diam.; spores 7-10 x 3.5-4.5 m; Bolivia C. complexum Filaments 4-8 p in diam. Filaments non-articulate; spermatia 7-9 x 15 /xj paraphyses cla- vate; Brasil C. dialeptizum Filaments articulate; spermatia not reported. Cells 2.5-4 times as long as broad, paraphyses obovoid; Brasil C. pannosum Cells 1.5-3 times as long as broad, sterile; Brasil C depressum Filament size not given, rugulose, articulate-branched, coalesced into fascicles; Venezuela C. Tuckermani Coenogonium Leprieurii NyL, Ann. Sci. Nat. Bot. IV, 16: 89. 1862. C. Linkii var. Leprieurii Mont., Ann. Sci. Nat. Bot. Ill, 16: 47. 1851. Holocoenis Leprieurii Clements, Genera of fungi, 174. 1909. Type: Guayane Frangaise, Leprieur, sine no. Algae loosely woven, yellowish, 11-16 (x in diameter. Apo- thecia plane or convex, excluding the white margin, disc livid from the first; asci about 20 y. in diameter; spores oblong to fusiform or ellipsoidal, unicellular, 6-10x2.5-4 (x; paraphyses slender, apices clavate. I have not seen this species from Costa Rica but Dodge & Nevermann 7399, from jungle at Castilla farm, 20 m., Lim6n , seems to be a variety of this species, developed but has a livid disc, the mar The fungus is and white, and the alga is of much smaller diameter. The exciple is composed of isodiametric cells with colored walls. It appears to be the same fungus attacking a much smaller alga, and should be referred to Patinella. Coenogonium subvirescens NyL, Flora 57: 72. 1874. C. Leprieur 16:89. 1862. NyL, Ann. Sci. Nat Brasil, Amazonas, Rio Negro, Spruce 28, is type of the 398 ANNALS OF THE MISSOURI BOTANICAL (JARDEN (Vol. 20 species, while C. Lcprieuru var. subvircsccns was described from French Guiana without collector. Algae somewhat interwoven but more or less parallel, 7-9 (a, forming a loose dimidiate to subeircular thallus depending upon position and shape of substrate, not or obscurely zonate, pale dark olive buff, cells ab evident ther cell walls thick, chloroplasts appearing somewhat dumb-bell with the pigment more or less massed ds of The Apothecia about 800 p. in diameter, tapering below to a short stipe about 250 n in diameter, composed of thick-walled pseudo- parenchyma with the outer 40 \x composed of a palisade of elongate cells, perpendicular to the surface; parathecium about 80 [x thick, homogeneous with the rest of the apothecium, the hypo- thecium very thin, filamentous, of cells with thinner walls, hymenium is about 80 [x tall and is composed of filiform l„„ physes with clavate tips and slender clavate asci. The asco- spores are not mature in my specimens but seem to be slender, 2-celled, probably about the size of most of the other spores in the genus except C. disjunctum. Limon: Cadiz, l)odg< : 452; Cast ilia farm, 20 m., Dodge & Nrvermann 7451; Monte Verde, A'. DanieUon 7H; Siquirres, 70-200 m., Dodge, Call & Thomas 5574; La Colombian*, 70 so m., Standley S6886. Cartago: R. I'ejivalle, 650-800 m., Dodge & Thomas 440S; El Mufieco, on R. Navarro, 1400-1600 m., Standley & J. Valerio 51055; C. Carpintera, 1500-1850 m., Standby 86696. Puntarenas: Osa, Golfo Dulce, Dodge 7 ',50. Guanacaste: Q. Serena noar Tilanin, 700 m., Standley ,fe J. Valerio 46242; Santa- maria, 720 860 Da., Dodge 7015. Coenogonium Linkii Ehrenl)erg ap. Nees ab Esenbeck, Horae l'liys. Berol. 120. 1820. C. controtrrsum Pers. ap. (Jaudich. in Kreycinet, Voy. Uranie Bot. 214. J<>zo Uiom Type: Brasil, Sta. Cntharina, Chamisso. The type of C. overswn came from the walls of the aqueduct of Corcovado, de Janeiro. Gai but 12 dimidiate to sub orbicular thallus, not or obscurely zonate, p buff, cells 40-00 [x long, septa very difficult to observe 1933] DODGE — LICHENS OF COSTA RICA. I 399 plasts small and scattered, outer cell walls thick, partially covered with anastomosing fungal hyphae. Apothecia warm buff, about 850 y. in diameter, tapering sharply to a short stipe about 240 [x in diameter and up to 300 ^ long, pseudoparenchymatous, without palisade layer at the sur- face; parathecium about 100 \i thick, homogeneous with the rest of the apothecium, the hypothecium very thin and soon evanes- cent; the hymenium about 80 u. tall, composed of filiform para- physes with swollen tips and cylindrical asci. The ascospores are not fully mature in my specimens but probably the common size for the genus. Cartago: C. Carpintera, 1800 m., K. Danielson 9. Alajuela: La Palma de S. Ram6n, 1100 m., Brenes 175. Guanacaste: El Silencio, near Tilaran, 750 m., Standley & J. Valerio U619. Coenogonium conferyoides Nyl., Flora 41 : 380. 1858. ? C. andinum Karsten ap. Nyl., Bot. Zeit. 20: 178. 1862. Type: Mexico, Orizaba, Fr. Mailer (Schimper Herb.). Type of C. andinum from Colombia, 2000 m., Lindig 2560, also Peru, Weddell, and southern Brasil, Guillemin. Since I have been unable to see the types, the interpretation of this species is difficult. In Ann. Sci. Nat. Bot. IV, 11: 242. 1859, Nylander amplified his very inadequate description, basing his emendations on a specimen from Tahiti, Lepine 14- When he monographed the genus in Ann. Sci. Nat. Bot. IV, 16: 91. 1862, he cited Lepine 14 from Tahiti first, adding Guadeloupe Island, Duchas- saing, Brasil, Gaudichaud, and Weddell, and Chile, Gay, not citing the original Mexican type. Since none of these descriptions was based on specimens having apothecia its application is further confused. Since the descriptions only state how it differs from C. Linkii, I have assumed that the thallus is more or with a larger alga than in that species. Nylander dimidiate also notes that a specimen from Colombia, Bogota, 2700 m., Lindig 887, has algal filaments 12-18 ;jl in diam. Unfortunately I have been unable to see any of the above-mentioned specimens. The following description is based on Costa Rican specimens. Algae somewhat interwoven but more or less parallel, 16-20 (-28) \l in diameter, forming a soft dimidiate or suborbicular thallus, not zonate, pale to deep olive buff, cells about 60-100 \l [Vol. 20 400 ANNALS OF THE MISSOURI BOTANICAL GARDEN long, septa plainly visible, chloroplasts rather indistinct in these specimens, often alternate cells more or less collapsing, giving a somewhat moniliform appearance to old filaments, outer walls thick, not densely covered with hyphae. The collapsed cells sometimes have the coloring matter collected at the ends, giving the dumb-bell appearance to the chloroplast as in C. subvirescens . Apothccia warm buff, 560 |x in diameter, patelliform, with stipe about 100 \i in diameter and 300 u, long, pseudoparenchymat- ous, of very thick-walled cells which tend to arrange their long diameters perpendicular to the surface of the apothecium, but not forming a definite palisade as in C. subvirescens; the para- thecium 40-50 [x thick, homogeneous with the rest of the apo- thecium, the hypothecium 20-30 [x thick of interwoven hyphae, the hymenium about 00 u. tall, composed of filiform paraphyses with spherical tips and slender cylindrical asci about 5 (x in di- ameter; ascospores distichous, 4-celled, 10-12x3 [x. [The asco- spores .ire immature and so closely packed in the ascus that I am not certain of size and septation, as I have been unable to free spores from the ascus. | Limon: Waldeck, Dodge 7/fiS. Cartago?: Morpho Valley, 1420 m., A'. DamUUonBl (This locality is uncertain and Danielson's barometric readings are apt to be a little high, but it probably in the vicinity of the upper Reventaz6n below Cartago, where Lepidoptera of the genus Morpho are not uncommon.] COENOCONIUM HETEROTRICHUM Mull. Arg., Bull. SoC. R. Bot. Belg. 32: 162. 1893. Type: Costa Rica, San Marcos de Dota, Tomiuz 6115. Thallus pulvinate, cespitose-effused, deep olive buff to dark olive, algae somewhat interwoven, dimorphic, larger filaments 20-25 tx in diameter, cells about 40 (x long, chloroplasts disciform, scattered; smaller filaments submoniliform, the cells of smaller diameter being about 7-8 u, and 12 [x long, those of larger diameter being about 10-12 tx and of about the same lemrth. br Fungus unknown. In Brenes c 29a the fungal hyphae are San Jose: S. MarcOfl de Dota, 1200 m., Tomiuz HI 15, type. Alajuela: La PalmadeS. Ramon, 12. , limusSOa. Coenocjonium interponendum Nyl. ap. Polakowsky, Jour Bot. Brit. & For. 15: 225. 1877. 1933] DODGE — LICHENS OF COSTA RICA. I 401 Type: Costa Rica, Cartago, Angostura, H. Polakowsky 496. Algae loosely interwoven, forming an adnate irregular thallus, deep olive buff to citrine drab, cells about 17-20 [x in diameter, 30-50 (t long, pigment quite uniformly dispersed throughout the cell, yellowish green. All my Costa Rican material is sterile. Nylander states that the spores are fusiform, 2-celled, 8 x 2.5 tx. Lim6n: Carmen, Dodge 7449. Cartago: Angostura, H. Polakowsky 496, type. Alajuela: La Palma de S. Ram6n, 1250 m., Brenes, 53a, 116, 399. Q 804 # &* Coenogonium interpositum Nyl., Ann. Sci. Nat. Bot. IV, 16:91. 1862. Type: Borbonia, Lepervanche, Mezieres; Louisiana, Hale. Thallus 20-28 ;x in diameter and 60-88 \l long, chl the shaoe of a dumb-bell with the pigment at the ends of the cells. [Apothecia pale carneo-luteous ; spores oblong, simple, 6-10 x 3 [x; paraphyses slender or medium.] As all my Costa Rican material is sterile the reference of it to this species is doubtful. It quite possibly belongs with C. disjunction Nyl., based on specimens from Martinique and Cuba, Wright 170, which reaches a slightly larger diameter of algal filament and much larger 2-celled spores. Both species are said to resemble C. confervoides and perhaps should be placed in the C. confervoides group. Cartago: C. Carpintera, 1560-1700 m., Dodge & Thomas 7920. Alajuela: La Palma de S. Ram6n, 1100-1250 m., Brenes 53, 115. Coenogonium interplexum Nyl., Ann. Sci. Nat. Bot. IV, 16:92, pi. 12, f. 20, 21. 1862. Type: Colombia, 2200 m., Lindig 2561. Thallus loosely interwoven, intricate, algal cells 12-16 [>. in diameter and 30 [l long, chloroplasts more or less disciform, scattered. [Apothecia fleshy-yellow, plane, 1 mm. in diameter, margin waxy, fleshy to white; spores short-fusiform, 2-celled, 8-10 x 3 \i; paraphyses medium, apex clavate.] Alajuela: Piedades de S. Ram6n, 900 m., Brenes 408. Puntarenas: Boruca, 560 m., Tonduz 6114. 402 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 20 Coenooonium pannosum Mull. Arg., Flora 64: 234. 1881. Type: Brasil, Sfio Paulo, Apiahy, Puiggari 10 J6. Thallus loosely interwoven, dark olive buff to citrine drab, more or less adnate to substratum, algal cells 0-8 [i in diameter, 28-32 [i long, pigment migrating toward the ends of the cells. [Apothecia 350-500 \i in diameter, pale or orange white, plane, with paler and thinner margin, finally immarginate and slightly convex, becoming more flesh color; hymenium hyaline, para- physes very slender with ob ovoid head about three times as thick as the stalk; asci slender, cylindrical, 8-spored; spores 2- locular, 6-7 x 2 n, fusiform with acute ends.] Since my Costa Rican material is sterile its reference here is uncertain and it may be only very young algae of C. subvirescens Nyl., with which it agrees in general appearance, but there is no trace of the formation of a dimidiate thallus. Guanacaste: II. Q. Azul, on slope of V. Tenorio, 500 000 m., Dodge & Thomas 665Jf. Coenogonium DEPRESSUM Mull. Arg., Flora 64: 525. 1881. Type: Brasil, Sao Paulo, Apiahy, Puiggari 10M+. Thallus adnate, pale olive buff, filaments short, flexuous, depressed, subintricate; cells 4-8(-13) y. in diameter, 16-20 (jl long, somewhat inflated in the middle but scarcely enough to give a moniliform appearance, chloroplasts no longer distinct. As I have not compared the Costa Rican material with the type I cannot be certain of the identity. Cartage): alpinr region of Frazil, f)rxted. Coenocjonium implexhm Nyl., Ann. Sci. Nat. Bot. IV, 16: 92. 1862. Type: Australia, Victoria, Jarvin, Ferd. Miiller. [Similar to C. interplexum but algal cells a little smaller, 10-13 (a in diameter, spores a little larger, 8-11 x 3.5-4.5 (i, paraphyses thicker.] The following collection has been referred here by Miiller Argau, but 1 have not studied it. Puntaronas: Horuca, 560 m., Tonduz 6113. Ephebaceae Thallus dwarf fruticose, branched, more or less filiform, without rhizinae, crustose or small squamose, with Scytone?na 1933] DODGE — LICHENS OF COSTA RICA. I 403 or Stigonema. Apothecia small, often with very small puncti- form disc, scarcely visible; paraphyses well developed or absent; asci 8-spored; spores hyaline, 1-2-celled. The individuals of this family are so small that they are easily overlooked both in collecting and in sorting material, and so the few species here reported are probably not representative of this group in Costa Rica. KEY TO TROPICAL AMERICAN GENERA OF EPHEBACEAE Thallus crustose to small squamose, homoeomeric; Brasil Pterigyopsis atra Thallus dwarf fruticose, dark, thickly branched. Apothecia sunken in swellings of the thallus, single or gregarious; spores unicellular; paraphyses present; Brasil Ephebeia Apothecia sessile on the thallus, either lateral or terminal. Thallus without pseudoparenchymatous cortex or medulla; paraphyses filiform, simple; asci 8-spored; spores ovoid or spherical, unicellular Thallus with pseudoparenchymatous cortex and medulla. Thermutis Spores unicellular Leptogidium byssoides Spores 2-celled Polychidium THERMUTIS E. Fr. Thermutis E. Fr., Syst. Orb, Veg. 1 : 392. 1825. Gonionema NyL, Mem. Soc. Sci. Nat. Cherbourg 3: 163- 1855. Thallus dwarf fruticose, thickly branched and filiform, without rhizinae, Scytonema present with the hyphae running in the gelified sheath. Apothecia small, lateral, saucer-shaped to almost spherical; parathecium often highly developed and thick; hypothecium light-colored ; paraphyses unbranched, filiform, tips not swollen; asci clavate, thin-walled, 8-spored; spores hyaline, ellipsoidal, elongate, unicellular, thin-walled. Sperma- gonia lateral or terminal, sessile, more or less spherical; spermatia small, ovoid, or elongate. Thermutis velutina (Ach.) Fw., Linnaea 23: 170. 1850. Lichen velutinus Ach., Lichenog. Suec. Prodr. 218. 1798. Gonionema velutina NyL, Act. Soc. Linn. Bordeaux 21: 262. 1856. This species has been reported by Muller Argau from the following localities in Costa Rica. The specimens were not studied while I was in Geneve. San Jos6: San Marcos de Dota, 1200 m., Tonduz 5378. Guanacaste: Boruca, 560 m., Tonduz 5371; Terraba, Tonduz 5372. 404 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 20 A small sterile specimen from Guanacaste: near Tilar£n, 500- G90 m., Dodge & Thomas 8081, may belong in Polychidium but the specimen is not definitely determinable as to genus. COLLEMACEAE Thallus gelified, (Tustose to foliose or dwarf fruticose, with or without rhizoids, sometimes umbilicate, homoeomerous with Nostoc. Apothecia from sunken almost perithecia to sessile apothecia usually with an amphilhecium, occasionally with parathecium; paraphyses simple; asci 8-spored; spores hyaline, spherical to acicular, straight or twisted, 1-celled to muriform, usually with a thin wall (except in Physma and section Lem- phospora of Lempholemma) . This family is widely distributed in the temperate and tropical zones, the more highly developed members occurring in the latter. 1 1 is possible that some of the smaller and less conspicuous genera have been overlooked or have been included in the crustose material in the preliminary sorting. So far only four genera have been found in Costa Rica. KEY TO TROPICAL AMERICAN GKXERA OF COLLEMACEAE Spores unicellular. Thallus crustose, not gelified, parathecium present; spores ellipsoidal, thin-walled. Brasil Leprocollema americanum Porto Rico L. Finkii Thallus squamulose, gelified; spores spherical, thin-walled hut surrounded by a thick gelif ied sheath. Thallus not corticate Lemphol Thallus corticate, wholly pseudoparenchymatous; Juan Fernandez. . . . t in ma Lemmop.sis polyschulioides Thallus foliose, gelified; spores ellipsoidal to fusiform, thick-walled, surrounded by a gelified sheath Physma Spores phragmospores or dictyospores. Cortex not developed; apothecia with paratheeia only Collcma Cortex of pseudoparenchyma; apothecia with amphithecia (paratheeia often also present) Leptogium LEMPHOLEMMA K6rb. Lkmimiolemma Korber, Syst. Lich. Germ. 400. 1855. Type species: Lempholemma com pactum Korber. Thallus from verrucose, squamulose, dwarf fruticose to foliose in our species, gelified when moist, clothed with rhizinae below, 1933] DODGE — LICHENS OF COSTA RICA. I 405 homoeomeric, without cortex, with Nostoc. Apothecia superfi- cial or terminal, mostly sunken in the thallus (not in our species) ; amphithecium either with or without cortex; parathecium color- less, urceolate; hypothecium colorless; paraphyses filiform, simple; asci 8-spored, clavate, often twisted below; spores hyaline, fusiform, ellipsoidal-ovoid or spherical, smooth. The genus has been divided into eight sections of which only one section Lemphospora is tropical. This section is characterized as having apothecia without cortex, spores more or less spherical with a thick gelified sheath. Lempholemma Dussii (Vainio) Zahlbr., Cat. Lich. Univ. 3: 23. 1925 [Collema Dussii Vainio, Ann. Acad. Sci. Fenn. A6 7 : 114. 1915] has been described from Guadeloupe in the Antilles. Lempholemma (Lemphospora) oblique-peltatum (Eschw. ap. Martius) Dodge, comb. nov. Collema oblique-peltatum Eschw. ap. Martius, Icon. PI. Cryptog. Fasc. 2: 27, pi 11, f. 2 [between 1828 and 1833]. l Type: Brasil, near Para, Martius. Thallus foliose, up to 5 cm. in diameter, drying grayish olive to dark olive gray, margin thick, broadly subdichotomously lobed, lobes up to 3 mm. broad and 2-3 mm. long, smooth, surface minutely pitted although appearing practically smooth to the naked eye, without soredia or isidia; thallus 200-400 n thick, homoeomerous, without cortex, but provided with a thin layer of tangled rhizinae below. Apothecia subsessile, often appearing obliquely attached to the thallus in section, up to 2.5 mm. in diameter, margin of the same color as the thallus, with periclinal folds and wrinkles, disc tawny to mars brown; am (d ecium wrinkles), homogeneous with the thallus, 200-300 [i thick but extending beyond the parathecium as much as 800 p; para- thecium pseudoparenchymatous, highly developed, about 100 [jl thick below, thinning out above to about 50 (x; hypotl about 40 [i thick, of highly gelified hyphae; thecium about 100 [l 1 1 have been unable to locate the date of the second fascicle of this work. The cover of the copy in the Missouri Botanical Garden gives the pages of text and plates in each fascicle but only the dates 1828-1834. Martius in his 'Flora Brasiliensis' 1 : 233-234. 1833, quotes from the 'Icones' citing page, hence it must have been issued before that date. In this work he placed it in the subgenus Enchylium. [Vol. 20 406 ANNALS OF THE MISSOURI BOTANICAL (5AHDEN herb tall; paraphyees about 1-2 [i in diameter, clavate above, septate, cells 4-0 [i long, with walls highly gelified so that they appear as slender rows of cells imbedded in a gel; asci clavate-cylindrical, about SOx 16 n, containing 8 monostichous spores; ascospores broadly ellipsoidal, 12 x S y. while still in the ascus, with a thick gelified wall. While I have not studied the type specimen of this species, a careful study of the description and the figures shows that it belongs in LemphoUmma section Lemphospora rather than in Collevia. Judging from determinations observed in various iria, this species seems to have been confused with Lepto- gium rcsiculosum {L. bullatum), L. tremel I aides , and L . foveolatum , although the abundant development of rhizoids below should have separated it. While it is minutely scrobiculate-foveolate (visible under hand-lens), it is not conspicuously so to the naked eye as in L. foveolatum. Besides my Costa Rican material I have also seen it from Eustis, Lake Co., Florida, Nash 2024, determined by Eckfeldt as L. foveolatum (sic) Nyl. < Limmi: Waldeck, Dodge 7404; near Siquirres, 70- 170 m., Dodge, Catt & Thomas 55S8, 8024; Hamburg, 66 m., Slandley & J. Valerio 48767. Guanacaste: H. Santamarfa, 720 m., Dodge A- Thomas 6898; near Tilaran, 670 m., Dodge it Thomas 6644, 8022. Pun tan-mis: near Coroial, 5-50 m., Dodge 8098; Puerto Jimenez, Brenes 839. Lempholemma (Lemphospora) dichotomum Dodge, sp. nov. Type: Costa Rica, Guanacaste, H. Granadilla, Dodge & Thomas 6736. Thallus foliosus, ad 5 cm. diametro, 400-050 [x crassitudine metiens, griseo-olivaceus, margine crassiuscula, integra, lobatus, dichotomus, lobi 1-1.5 mm. lati, divergentes, superne longi- tudinaliter rugosus, sine sorediis isidiisque, inferne rhizinis nigris densissimis intertextis obsitus, homoeomerus, nostocaceus, sine strato corticali. Apothecium peltatum, basi constrictum, ad 2.5 mm. diametro metiens, margine concolori, rugosa, disco fulvo castaneove; amphithecium sectione lobatum, homogeneum, 150-300 [l crassitudine inferne, sed ad 500 ^ superne et ultra inarginem parathecii; parathecium pseudoparenchymaticum, ad 160 \i crassitudine inferne, attenuatum ad 6-9-septate; Brasil v. brasiliense Spores 36-50 x7 p, only 5-septate C. leucocarpum Apothecia not white-pruinose. Spores 48-66 x 6-8 mJ Colombia C. implication Spores 38-50 x 6-8 lx; Chile C. chilenum Spores 40-45 x 5-6.5 lx) apothecia sessile, margins turgid; Mex- ico C. turgidulum Spores 30-37 x 2.5-3 lx; apothecial margins thin, blackening; Brasil C. baculiferum Collemodiopsis Vainio, fitude Lich. Bresil 1 : 234. 1890, Type species: Collema nigrescens (Huds.) DC. [Vol. 20 410 ANNALS OF THE MISSOURI HOTANICAL GARDEN Spores slender, several-celled phragmospores, never muriform, apothecia with a pseudoparenchymatous cortex. This subgenus is typically northern in its distribution. Only C. Ihcrm'uneri Hue from Guadeloupe has been described previ- ously from the American tropics. Collema (Collemodiopsis) Granadillae Dodge, sp. nov. Type: Costa Rica, Guanacaste, H. Granadilla, Dodge & Thomas 6576. Thallus irregulariter subpinnatifide lobatus, lobis tenuibus, ad 1 mm. latis, viridi-nigricans, superne elevate )-rugosus verruco- susque ad subisidiosus, inferne reticulatim elevato-rugosus, cinerascens, ad 400 [l crassitudine, decorticatus, rhizinis desti- tutus. Apothecium planum, peltatum, basi constrictum, 0.5- 1.0 mm. diametro metiens, omnino nigrum, margine verrucosa; amphit liecium 1 50-250 \l crassitudine, pseudoparenchymatice subrectis; parathe pseudoparenchym crassitudine inferne, attenuatum ad 25-35 [x superne; hypothecium hyphis tenuibus dense contextum, 25-30 [x crassitudine; thecium 150- 170 [a altitudine, paraphyses arete cohaerentes septati, cylindrici, filiformes, ad 1 ;x diametro metientes, clavato-capitati; ascosporae octonae, polystichae, rectae, fusiformes vel aciculares, 5-septatae, 37-42 x 4 5 (jl. Thallus irregularly subpinnately lobed, lobes slender, about 1 mm. broad, dark greenish black, with elevated folds and wrinkles above with masses of small subisidiose warts, below reticulately deeply scrobiculate wrinkled, more or less ashy, about 400 [x thick, without cortex or rhizoids. Apothecium peltate with constricted base, plane, about 0.5-1 mm. in diameter, wholly black, margin verrucose; amphithecium 150-250 [l thick, pseudoparenchym a below filaments of Nostoc woven; parathecium of large-celled pseudoparenchyma, about 100-120 (jl thick below r , thinning above at the margin to 25-35 [x; hypothecium of densely woven slender hyphae, 25-30 [x thick; thecium 150-170 [x tall; paraphyses closely adherent, septate, cylindrical, filiform, about 1 [x in diameter, clavate, capitate at the apex; ascospores 8 per ascus, polystichous, straight, fusiform or acicular, 6-celled, 37-42 x 4-5 tx. 1933] DODGE — LICHENS OF COSTA RICA. I 411 Standley 640-660 Collema leucopepla (Tuck.) Schneider, Guide Study Lich., 181. 1898. Collema nigrescens var. leucopepla Tuck., Syn. N. Am. Lich. 1 : 148. 1882. Type: not stated in original description, based on material from South Carolina, Georgia, Florida, Alabama, and Louisiana. It is probable that material collected in Puntarenas, Boruca, 560 m., Tonduz 5373, and determined by Miiller Argau as C. nigrescens var. caesium Ach., belongs here but I did not have time to study it critically while I was in Geneve. This species of the lower Atlantic and Gulf Coastal Plain in the United States has also been reported by Vainio from Mexico. DICOLLEMA (Clements) Dodge, n. subgenus. Dicollema Clements, Gen. Fung. 74. 1909. Type species: Collema pycnocarpum Nyl. Thallus narrowly laciniate or incised, spores clavate to fusiform, typically 2-celled, rarely 4-celled. The material referred to the type species of this genus by different authors is quite variable and the whole group needs a thorough revision. Pending such a study, I am proposing several varieties of C. pycnocarpum Nyl. to cover the more conspicuous variations which seem to have separate geographical ranges, those with the smaller spore sizes being rather more southern in their distribution. Collema pycnocarpum Nyl., Syn. Meth. Lich. 1: 115. 1858. Type: United States [on journey from New York to Philadel- phia], More. Thallus dark or pale green, medium in size, granulate-nodose. Apothecia rufous, crowded, almost contiguous, at first plane, then somewhat convex, quite small, 0.5 mm. or a little larger; spores 8 per ascus, oblong or oblong-ellipsoid, simple or uniseptate, 14-17 x 4-5 n. Northern Atlantic Coastal Plain. Var. Minarum Dodge, var. nov. Collema pycnocarpum Vainio, fitude Lich. Br6sil 1 : 238. 1890. Type: Brasil, Minas Geraes, Sitio, 1000 m., Vainio 734- 412 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 20 Sporae oblongae, ellipsoideae vel fusiformes, 1-septatae, 6-9 x 3 (x. Var. Malmei Dodge, var. nov. C. pi/rnocarpum Malme, Ark. f. Hot. 19 8 : 7. 1924. Type: Brasil, Rio Grande do Sul, Canoas near Porto Alegre, Malme 586, Sporae oblongo-ellipsoideae vel ellipsoideae, uniseptatae, 10- 14 x 3-4 |a. Var. crassiusculum (Malme) Dodge, comb. nov. Forma crassiusculum Malme, Ark. f. Bot. 19 s : 7. 1924. Collema pi/cnocarpum Nyl., Acta Soc. Sci. Fenn. 7: 428. 1803. Type: Brasil, JVlatto Grosso, Corumba, Malme. Sporae ellipsoideae, utroque apice obtusae, raro acutae vel altero obtusae, altero acutae, 10-15.5 x 4-5.5 [x. The specimens cited by Nylander from Colombia, Bogota, 2000 m., Ldndig 2872, have spores 10-14 x 4.5-5.5 ji. Collkma cyrtaspis Tuck., Proc. Amer. Acad. Arts & Sci. 5:387. 1802. Type: no type mentioned in the original description. Thallus deeply and irregularly lobed, verrucose, deep blue green, darkening on drying, verrucae much larger than in C. pyenocarpum, about 400 |x thick; algal layer about 100 |x thick, composed of coiled and tangled filaments of Nostoc imbedded in a gel, cells spherical, about 4 [x in diameter; medulla gelified, traversed by very loosely tangled hyphae about 2-3 |x in diameter and occasional straight filaments of Nostoc. Apothecia immersed in the verrucae when young, becoming plane or even convex, up to 2 mm. in diameter, margin slightly verrucose when young, practically disappearing as the apothecium becomes increasingly complex; amphithecium about 160 ;x broad, of the same texture as the thallus; parathecium lacking; hypothecium filamentous, quite highly developed, about 40 jx thick, not conspicuously thinner toward the margins, hyphae 2-3 [x in diameter, very loosely woven; thecium about 80 fx tall; paraphyses filamentous, clavate, expanded above, imbedded in a gel; asci clavate, about 40x8 [i, 8-spored; ascospores subfusiform, 2-4-locular, 10-25 x 3-7 [x [10-20 x 4-4.5 (x in Costa Rican specimens]. This species is widely distributed in the southern Atlantic 1933] DODGE — LICHENS OF COSTA RICA. I 413 and Gulf Coastal Plain in the United States, extending up the Mississippi Valley to Iowa. Guanacaste: Liberia, 100 m., Dodge & Thomas 8016, 8027. Synechoblastus (Trevis.) Vainio, Etude Lich. Br£sil 1 : 234. 1890. Lathargium S. F. Gray, Nat. Arrang. Brit. PL 1: 399. 1821, p.p. Synechoblastus Trevis., Caratt. Tre Nuov. Gen. Coll. 3. 1853, p. p. Type species: as subgenus based on C. glaucophthalmum Nyl. Apothecia without pseudoparenchymatous cortex, spores fusi- form to more or less acicular, many-celled, mostly over 30 [i long. This subgenus is typically northern. Only two species are known from Costa Rica, where they are temperate species at elevations between 1000 and 1800 m., and are also found in the highlands of Colombia at somewhat greater altitudes. Collema glaucophthalmum Nyl., Syn. Meth. Lich. 1: 114, 115. 1858. Type: Mexico, Orizaba, Fr. Midler. Thallus olivaceous-fuscous, medium size, expanded, more or less fenestrate and dissected, scrobiculate, and often granuliferous. Apothecia glauco-lilac colored, plane and somewhat concave, margin prominent, thin; ascospores 77-92 x 6-7 \i. Var. granatense Hue, Jour, de Bot. [Morot] 20: 12. 1906. Collema glaucophthalmum Nyl., Acta Soc. Sci. Fenn. 7: 428. 1863, non loco alio. Type: Colombia, Choachf, 2600 m., Lindig 813. Thallus dusky yellowish green to almost black, irregularly lobed and fenestrate, smooth or somewhat scrobiculate, coarsely wrinkled and subverrucose, especially toward the tips, margins thick and rounded, 800-1000 \i thick, homoeomerous, of loosely tangled filaments of Nostoc y cells 5-6 x 2.5-3 [i, ellipsoidal. Apothecia peltate, constricted at the base, up to 3 mm. in di- ameter, margin at first prominent, smooth, becoming thinner, less prominent and verrucose at maturity; disc chalky white to vinaceous-russet ; amphithecium about 160 \l thick, homogeneous with the thallus; parathecium pseudoparenchymatous, 40 [x thick, thinning to 20 \i or even disappearing above at the margin ; I Vol. 20 414 ANNALS OF THE MISSOURI BOTANICAL GARDEN hypotheeium filamentous, of large very densely interwoven hyphae about 40 \l thick; thecium about 120 ^ tall; paraphyses filiform, about 1 y. in diameter, not swollen above; asci clavate, about 20 [x in diameter, 8-spored; ascospores polystichous, fusiform to acicular, 55-74 x 5-7 (x. The whiteness of the apothecial disc is very variable on the same thallus, in general being more pronounced on young and rapidly growing apothecia and gradually disappearing on older and more exposed ones. It seems to be a semi-crystalline deposit which slowly dissolves away, disclosing the vinaceous- russet disc formed by the discoloration of the upper portion of the gel surrounding the paraphyses. This variety has previously been reported only from Colombia. Cartago: near R. Birrfs above Santiago, 920-1340 m., Dodge 8011, Dodge A Thomas 8015; Carpintera, 1700 in., A'. DunicUon 102. San .lost' : St.i. Maria de Dota, 1500-1800 m., Slandley & J. Valcrio U150. Collema implicatum Nyl., Acta Soc. Sci. Fenn. 7: 428. 1863. Type: Colombia, Villeia, 1100 m., Lindig 749; Cundinamarca, Bogota, 2400-2600 m., Lindig. Thallus dark greenish black, broadly lobed and fenestrate, I smooth near margins, becoming deeply reticulate-scrobiculate in older portions of the thallus, verrucae scattered, more elevated, often subisidioid, margins thin and semipellucid, homoeomerous, of loosely tangled filaments of Nosloc, cells 4x5 [x, heterocysts 5 x 8 [k, much more closely tangled in an outer zone about 150 [i thick, Apothecia peltate, constricted at the base, up to 4 mm. in diameter, margin at first prominent, verrucose, becoming thinner, less prominent, scarcely visible as the apothecium becomes expanded and convex, disc cameo-brown, without pruina in young apothecia; amphithecium about 100 (x thick, homogeneous with the thallus; parathecium pseudoparenchy- matous, about 80 \l thick below, thinning out to a single layer of thick-walled cells 6 [/, wide above at the margin; hypotheeium of densely woven hyphae about 20 \i thick; thecium about 100 [l tall; paraphyses about 2 \i in diameter, with clavate brown tips; asci clavate, 8-spored, 12-10 [i in diameter; ascospores poly- stichous, fusiform to acicular, 48-06 x 0-8 n, 7-9-septate. This species is very closely related to C. glaucophthalmum 1933] DODGE — LICHENS OF COSTA RICA. I 415 and has been reduced to synonymy by some authors. It differs from the previous species in several minor details of proportion of measurements and in the lack of a well-developed pruina. It is to be hoped that some one will be able to study the two forms in the field to determine the constancy of this character. In a series of specimens one can find considerable variation in the amount of pruina in C. glaucophthalmum. It should be noted that the two species occupy the same areas in Costa Rica. 140-1180 1700 m., K. Danielson 101c. _ r s has not been found in Costa Rica but it is closely related to this group. Collema (Synechoblastus) Ramboi Dodge, sp. nov. Type: Brasil, Rio Grande do Sul, Porto Alegre, B. Rambo 74- Thallus parvus, adscendens, obscure viridi-nigricans, angusti- lobatus, fenestratus clathratusque, verrucis elevatis isidioideis, marginibus plus minusve integris, tenuibus, pellucidis, 130-140 [x crassitudine, homoeomerus, filamentis nostocaceis laxe implexis, in zona exteriori 40 [l crassitudine dense contextis, cellulis ad 3x6 (jl. Apothecium peltatum, basi constrictum, planum, 0.5-0.6 (- 1.0) mm. diametro, margine tenui, verrucosa sub- crenulatave, disco obscure castaneo nigroque; amphithecium 20 p, crassitudine, cum thallohomogeneum; parathecium deest; hypo- thecium 20 [x crassitudine, hyphis tenuibus dense contextum; thecium 160 p altitudine; paraphyses filiformes, ad 1 ^ diametro, apicibus inflatis; asci clavati, 120x8-10 p; ascosporae octonae, polystichae, aciculares, immaturae, plus quam 100 [l longitudine, multiloculares. Thallus small, elevated, dark greenish black, narrowly lobed and fenestrate, verrucae elevated and isidioid, margins more or less smooth, thin, pellucid, 130-140 [i thick, homoeomerous, of loosely tangled filaments of Nostoc, cells about 3x6 n, much more closely tangled in an outer zone about 40 [l thick. Apothecia peltate, constricted at the base, 0.5-0.6 (- 1.0) mm. in diameter, plane, margin very thin, verrucose, subcrenulate, finally almost disappearing while the expanded apothecium remains plane, disc dark chestnut to black; amphithecium 20 \l broad, homo- geneous with the thallus ; parathecium absent ; hypothecium 20 (x 416 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 20 thick, of slender densely woven hyphae; thecium about 100 [i tall; paraphyses filiform, about 1 [i in diameter with greatly swollen tips, imbedded in a hymenial gel; asci clavate, 8-spored, 120 (jl long, 8-10 [x wide; ascospores polystichous, 8 per ascus, acicular, immature and very difficult to measure but somewhat more than 100 (x long, many-celled. LEPTOGIUM S. F. Gray Leptogium S. F. Gray, Nat. Arrang. Brit. PL 1: 400. 1821. Type species: Leptogium tremelloides S. F. Gray, excl. syn. Thallus foliose in the tropical species, gelified, below naked (covered with rhizinae in section Malloiiwn), with pseudo- parenchyma I ous cortex above and below, algae Nostoc. Apo- thecia sunken in the thallus at first, emerging and often short- stipitate; amphithecium often corticate with several layers of pseudoparenchyma below, usually of a single layer above; parathecium usually present and pseudoparenchymatous, often thinning out above at the margin, sometimes of large, more or less parallel hyphae; paraphyses simple and filiform, often with apex variously thickened; asci clavate to cylindrical, 8-spored; ascospores usually imbricately monostichous, occasionally distich- ous, hyaline, usually broadly fusiform with acute to acuminate ends (acicular in section LcpUxjioysis), usually muriform (except in Leplogiopsis), with thin walls. Four of the seven sections into which Leptogium is divided are found in tropical America, all four of them in Costa Rica. Lep- togiopxis differs from all the other sections in its acicular, never muriform spores, and perhaps should be treated as a separate genus, more or less related to section Syncchoblastus of Collcma. It is endemic in tropical America, with a single unidentifiable fragment yet found in Costa Rica. Mcdlotium, to which may also be referred the section Leptolobaria of Vainio based on L. callithamnium from Chile and the Antilles, has highly developed rhizinae below, and often more or less tomentum above, especially in the vicinity of the apothecia. It is widespread in its distribu- tion, being found in the temperate zones and evidently preferring the colder regions of the tropics, not occurring below about 1300 m., in Costa Rica not seen below 3000 m. DiplothaUus, with 1933] DODGE — LICHENS OF COSTA RICA. I 417 its two separate layers of thallus, each with its own cortex, connected by pillars of pseudoparenchyma which on drying cause punctate depressions in the thallus, is endemic in tropical America and confined to the temperate regions, in Costa Rica occurring between 1200 and 1700 m., coming down to lower levels in the mountain near H. Santamaria of the Cordillera de Tilaran in Guanacaste. Euleptogium, which contains the most species, is found from sea level to about 1800 m. in Costa Rica, although most of the species of this section have a narrower altitudinal range. The lowland species ascend the river valleys but mostly drop out at about 1000 m. These are usually wide- spread at low elevations, extending from the Atlantic and Gulf Coastal plains of the United States to southern Brasil and Paraguay. The species of the temperate region of Costa Rica are mostly confined to the mountains from southern Mexico to the northern Andes and the mountains of eastern Brasil. The morphology of the apothecium is not altogether clear and considerable confusion exists in the nomenclature of the various parts. In the following discussion, I have treated the tissue, usually well differentiated, underlying the hypothecium, as the parathecium, whether it extends to the surface of the thecium or not and whether or not it is pseudoparenchymatous. In a very few species the parathecium as thus defined thins out and disappears near the edge of the thecium, and the hyphae of the hypothecium extend up to the surface of the thecium. It is possible that some would prefer to regard the parathecium as formed of two layers, the lower of which is pseudoparenchymatous and the upper filamentous, with only the upper extending to the surface of the thecium. The amphithecium (thalline margin of earlier lichenologists) is essentially homogeneous and continuous with the thallus, although the algae are sometimes more densely tangled and in many species the cortical cells on the under side of the amphithecium (morphologically continuous with the upper surface of the thallus) or even the under side of the thallus below the apothecium, may proliferate, forming a pseudoparen- chyma of several layers of cells sometimes even thicker than the algal zone. [Vol. 20 418 ANNALS OF THE MISSOURI BOTANICAL GARDEN KEY TO THE TROPICAL AMERICAN SPECIES OF LEPTOGIUM Spores acicular, never muriform. Thallus retioulate-scrohioulate; French Guiana. [Spores 0-celled, 35 x 10 li, teste Leighton, in specimen from Amazonas, Brasil] L. reticulatum Thallus not reticulate-scrobiculate. Spores IS ix long, 8-10-celled; Florida L.fusisporum (Tuck.) Dodge Spores 60 60 x 7 S li } 10-12-celled; Mexico L. adpressum Spores Of) -85 x 3.5-5 /i, 10 12-celled; Rrasil L. megapotatnicum Spores muriform. Thallus of two lamellae, each with its own cortex, attached to each other by columns of tissue DIPLOTHALLUS A single species L. diaphanum Thallus of a single lamella, not as above. Thallus with a thick tomenium of rhi/inae below MALLOTIUM Apotheeia on lower surface, lobes of thallus broad, imbricate. . . .L. rcsupinans Apotheeia on upper surface. Upper surface of thallus tomentosc, at least near apotheeia. Margins of sinuses in rolled, lobes medium, sinuate; apothecial margins microphylline L. inflcrum Margins of thallus isidiose dissected. Upper surface of thallus subglabrous and more or less granular- papillate L. papillosum Upper surface of the thallus arachnoid-tomentose, lobes minutely dissected, not otherwise isidiose; ( !hile.L. callithamnium Upper surface of thallus glabrous except in the vicinity of the apotheeia; apothecial margin isidiose, scattered groups of isidia sometimes on the upper surface of the thallus. L. inflexum v. isidiosulum Thallus corticate with a single layer of cells on both surfaces, without rhizinae EULEPTOGIUM Thallus scrobiculate. Margin of thallus papillate-denticulate, olivaceous, thallus deeply reticulate-scrobiculate; paratheeium filamentous; spores 22- 21 x 12-14 m L. olivaeeum Margin of thallus smooth, thallus plumbeous. Thallus deeply reticulate-scrobiculate; spores 30-40 x 12-16 \x. L. foveolatum Thallus shallowly scrobiculate; spores 20-30 x 8- 10 m- • -L. microsticlum Thallus variously wrinkled on drying, but not scrobiculate, usually nearly smooth when moist. Apotheeia on tips of long swollen processes of the thallus. Thallus lobes narrow, dichotomously branched, wrinkles ir- regular L. stipitatum Thallus lobes broader, not conspicuously dichotomously branched, wrinkles predominantly longitudinal or radial, secondary wrinkles mostly perielinal to margins. Apotheeia radially wrinkled t<> smooth; spores 23-32 x 10 13 /x L. vesicidosum Apotheeia with perielinal wrinkles, foliolate or isidiose. Thallus and apotheeia not isidiose. 1933] DODGE — LICHENS OF COSTA RICA. I 419 Apothecia small, spores 22-30 x 10-12 p, margin wrinkled. Thallus lobes broad L. phyllocarpum Thallus lobes narrower, margins crisped v. campestre Apothecia rather large with more or less foliolate margins. v. macrocarpum Thallus and apothecia isidiose, often densely so; spores 30~40(-45) x 12-17 m L. coralloideum Apothecia marginal, sessile or on short solid stalks, very minute. Thallus dark green to black, exciple isidiose [spores 24-30 (-33) x 12-14 m, transverse septa 5, fide Malme]; Jamaica L. chloromelan Thallus ashy to plumbeous. Exciple isidiose, parathecium 150 m thick; thecium 150- 180 m tall L. marginellum Exciple verruculose, parathecium 100 m thick; thecium 120-140 n tall; Paraguay L. pilcomayense Exciple crenulate, parathecium 30 m thick; thecium 110- 130 m tall; Paraguay L. microcarpum Apothecia scattered on upper surface of the thallus. Thallus thick, 250 m or more, roughened below by more or less conical outgrowths but rhizinae absent. Smooth above or slightly wrinkled, lobes ascending, obo- vate; Mexico L. hypotrachynum Reticulately wrinkled above, lobes appressed, short, wide, margins sometimes crisped; Brasil L. mattogrossense Smooth or only slightly wrinkled below L. sessile Thallus thinner, seldom reaching 200 fx. Thallus more or less isidiose. Isidia blackening, cylindric; parathecium well developed. Spores 18-23x8-10 ^ transverse septa 3, rarely 5; Brasil and Paraguay L. pichneum Malme excl. syn. Spores 22-30 x 11-13 m, transverse septa 5 L. simplicius v. pichneoides Isidia concolorous, parathecium not well developed L. denticulatum Nyl. Thallus 100 m thick; apothecia not isidiose; spores 22-27 x 7-9 n, transverse septa usually 5; isidia coralloid branched or somewhat flattened; Brasil. L. austroamericanum (Malme) Dodge Thallus about 30 m thick; apothecial margins verrucose, spores 16-23 x 8-9 m, transverse septa 3; isidia often flattened and microphylline L. denticulatum Malme non Nyl. Isidia of hemispheric concolorous granules, thallus rigid, irregularly undulate, wrinkled toward the margin; Paraguay L. granulare Thallus not isidiose. Parathecium filamentous, or, if pseudoparenchymatous, very thin and inconspicuous. [Vol. 20 420 ANNALS OF THE MISSOV1U HOTANICAL GARDEN Cortex of amphithecium a single layer of cells; thallus 50-75 fi thick; apoihecia rarely 1 mm. in diam.; spores 15-11> x 6-8 n with 3 transverse septa; Brasil . L. Puiggarii Cortex of amphithecium a single layer of cells above, becoming thick, pseudoparenchymatous below; apoihecia I \\ mm. in diam. Thallus greenish black, 80-100 /* thick; spores 16-22 x 6-8 Mt with 3-5 transverse septa; Brasil. . .L. brasiliense Thallus ashy, plumbeous, or bluish. Thallus papulose and wrinkled above, opaque, 130-160 m thick; spores 20 30 x 7-12 M . .L. pulchellum Thallus minutely wrinkled above, opaque, 150- 170 n thick; spores 24-28 x 12 p L. Standleyi Thallus minutely wrinkled above, margins in- rolled; spores 38-46 x 16-18 M L. dimarphum Thallus smooth or minutely wrinkled on drying, pellucid, less than 100 m thick. Spores 22 28 x 8-10 p; thecium 120-150 M tall; thallus 80-100 M thick L. az Spores Is 22 x 10 12 M , with 3-5 transverse septa; thecium 90-120 \i tall; thallus 35- 60 m thick L. Tuckermani Spores 18-24 x 8-9 \x with 3 5 transverse septa; thecium 140-160 M tall; thallus 35-50 M thick; Brasil L. Schiffneri ure urn Parat hecium pseudoparenchymatous. Spores 35-40 x 16-20 ^; thallus plumbeous to greenish; amphi- thecium smooth; Bolivia L. luevius (Nyl.) Dodge Spores 30-40 x 12-17 p; thallus rather dark plumbeous to black. Laciniae 1-1.5 mm. broad, apothecia not over 1.5mm. in diam.; thecium 170 a* tall; paraphyses 1.5-2 ^ thick; Brasil L. Lafayrttcanum Laciniae up to 8 mm. broad, apothecia 1-2.5 mm. in diam.; thecium 190-220 M tall; paraphyses 2.5-3 m thick; Brasil. L . par h ych til u m Spores 25-35 x 12-15 m; apothecia 1 2.5 mm. in diam.; thecium 150-180 ^ tall; paraphyses 2-2.5 M thick; Brasil I j. vivaa u m Malme non Pers. ictus Spores 22-30 x 1113 /x; apothecia 1.5-2.2 mm. in diam.; thecium 150-170 M tall L. simplici Spores 18-34 x 9-15 mJ thallus dark green to plumbeous; amphi- thecium granulate to wrinkled. Thalline lobes ascendant, gy rose-plicate. .L. conchatum (Tuck.) Dodge (L. cMoromdan auct. non Sw.) Thalline lobes narrower, edges erect, crisped. L. BteUans (Tuck.) Dodge Dip loth allus Vainio, Etude Lich. Br&sil 1: 222. 1890. Thallus of two lamellae connected by pillars. Each lamella 1933] DODGE — LICHENS OF COSTA RICA. I 421 has a cortex of a single layer of isodiametric cells above and below, drying impressed-punctate. Rhizinae absent, spores muriform. Leptogium diaphanum (Sw.) Mont., Ann. Sci. Nat. Bot. Ill, 10: 134. 1848. Lichen diaphanum Sw., Nov. Gen. Sp. PL Prodr. 147. 1788. Parmelia diaphana Ach., Meth. Lich. 223. 1803. Collema diaphanum Ach., Lichenog. Univ. 654. 1810. Leptogium punctulatum Nyl. ap. Fournier, Mexic. PL 1: 1. 1872. Leptogium tremelloides var. impressopunctatum Tuck. ap. Williams, Amer. Nat. 29: 482. 1895. Type: Jamaica, Swartz, in Riksmuseet, Stockholm, carefully described by Malme, Ark. f. Bot. 19 8 : 27. 1924. Thallus mineral gray, lobes rounded, ascending, crowded, impressed-punctate, otherwise quite smooth, consisting of two layers, each about 40 \i thick, composed of a row of pseudo- parenchymatous cells about 8 ^ in diameter on each surface and an algal layer of Nostoc between, the layers connected by pseudoparenchymatous pillars whose contraction in drying form the depressions. Apothecia borne on the upper layer, peltate, constricted at the base, 1-1.5 mm. in diameter, margin light buff, smooth, disc chestnut; amphithecium corticate with thick pseudoparenchyma below, about 80 (x thick, thinning to two layers of cells above next the disc, algal layer about 30 (x thick below the thecium; parathecium absent; hypothecium of densely woven, large, thick- walled hyphae about 50 [x thick; thecium 120-130 [i tall; paraphyses filiform, 1-2 [x thick, with clavate tips, forming a brown epithecium imbedded in a gel ; asci clavate to cylindric, about 12-16 [x in diameter, thin- walled tips thickened and staining deep blue with iodine ; spores 8 per ascus, imbricately monostichous, fusiform, muriform, 16-25 x 7-8 (x, with 3-5 transverse septa. This species is found in the mountains from Mexico and the West Indies, Dominican Republic, and Jamaica (900-1200 m.) to Minas Geraes (1400-1500) m., Bolivia (1900 m.), and Peru (1700 m). In Costa Rica it is found from 1200 to 1700 m. and descends to 800 m. on the mountain back of the farmhouse at 422 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 20 H. Santiunurfa where the peculiar weather conditions enable many species to flourish below their normal altitudes. Ap- parently it needs high humidities as it is found mostly in localities of frequent and long-continued fogs (see p. 379). CartaRo: R. Birrfa above Santiago, 1220-1340 m., Dodge & Thomas 7938; Cartago, R. Toms R. 143; Carpintera, 1700 m., A'. Danideon 103. San Jose: Bta. Maria tie Dota, 1500-1800 m., Standby & J. Valeria 4320S. Heredia: C. Central do Zurquf, 1600-1700 m., Dodge, J. Valeria & Thomas Jf62J^. Guanacaste: II. Santamarfa, 720-850 m., Dodge, Jimenez & Thomas 7017. Mallotium Ach., Lich. Univ. (544. 1810. M alio Hum Gray, Nat. Arrang. Brit. PL 1: 399. 1821. Type: Collema saturninum (Dicks.) Ach. Thallus foliose, cortex of pseudoparenchyma above, below tonientose with rhizinae; spores muriform. This subgenus, considered as a separate genus by many authors, is cosmopolitan in distribution, mostly along cold foggy coasts or in mountains, being abundant in species in northern Europe and in Patagonia, Tierra del Fuego, and Ant- arctic Islands. Our tropical species seem confined to much higher elevations (above 1800 m. in Costa Rica) than the other members of Leptogium. It is quite distinct in appearance, often tonientose above, so that sterile specimens might be taken for Erioderma or Umbilicaria on macroscopic examination. Our two species, L. inflexum and L. papillosum, are both Mexican, the former extending to Peru, the latter not known south of Costa Rica. Leptogium papillosum (Bouly de Lesdain) Dodge, comb. nov. Leptogium Hildenbrandii var. papillosum Bouly de Lesdain, Lich. Mexique, 30. 1914. Type: Mexico, Michoacan, Puebla, H. Batan, G. Arsdne Brouard 1$12; Morelia, C. Azul, 0. Arsene Brouard 8999. Thallus mineral gray above, dark olive buff below, lobes subpinnatifid, with small, lacerate, almost isidioid margins, papillate-granulose, sometimes isidiose above, densely tonientose below, algal layer about 60 \l thick, corticate on each surface, with cells 5-6 \l in diameter, the cortex of the lower surface giving rise to a dense covering of rhizinae. Apothecia sub- marginal but too immature in our specimens to show details of structure well. 1933] DODGE — LICHENS OF COSTA RICA. I 423 San Jose: L. de la Chonta, n. e. Sta. Marfa de Dota, 2000-2100 m., Standley 42285; near Sta. Maria de Dota, 1500-1800 m., Standley 41643. Leptogium inflexum Nyl., Flora 41: 377. 1858; Syn. Meth. Lich. 1 : 132. 1858. Type: Mexico, Orizaba, Fr. Mutter. Thallus deep glaucous gray, dark olive buff below, lobes ir- regular, rounded, margins crisped and subascending, smooth above except in the vicinity of the apothecia, densely tomentose below, algal layer about 60 y. thick, corticate on both sides with two layers of cells about 8-10 \x in diameter, the lower surface giving rise to rhizinae, as also the outer cortical cells in the vicinity of the apothecia. Apothecia large, up to 4 mm. in diameter, margin more or less tomentose below, phyllophorous, concolorous with the thallus, disc orange rufous to auburn, edge of para- thecium lighter; amphithecium with pseudoparenchymatous cortex about 100 y. thick, thinning to about one or two cells thick above, algal layer of Nostoc about 80 \l thick; parathecium pseudoparenchymatous, about 20 y. thick below the hypothecium, expanding upward to thickness of 100 y. above; hypothecium 30 y. thick, of densely woven hyphae; thecium 240 y. tall; para- physes filiform, 1-2 y. in diameter, with swollen clavate tips forming a brown epithecium ; asci 20-25 y. in diameter, cylindrical ; ascospores monostichous, 8 per ascus, broad, fusiform, muriform, with 7 transverse septa, about 35-40 x 13-17 y.. Superficially the thallus suggests the texture of L. tremelloides , while the apothecia might easily be mistaken for L. phyllocarpum, although the rhizinae below should be easily observed. Cartago: F. Volcan de Turrialba, 2000-2400 m., Standley 34966, 35174- San Jos6: Quebradillas, 7 km. n. Sta. Marfa de Dota, Standley 42899; C. de las Vueltas, 2700-3000 m., Standley & J. Valerio 43823a. Var. isidiosulum Nyl., Acta Soc. Sci. Fenn. 7: 429. 1863; Ann. Sci. Nat. Bot. IV, 19: 289. 1863. Type: Colombia, Paramo Choachi, 3600 m., Lindig. This variety differs in the thallus being partly isidio-furfu- raceous, especially the margins of the lobes; apothecial margin isidiose instead of phyllophorous, subnude below ; spores 36-44 x 18-25 (x. This variety would seem to differ from L. papillosum in that the isidia are in scattered groups or marginal, much more highly 424 ANNALS OF THE MISSOURI BOTANICAL GARDEN |Vol 20 developed and more denuded below. A single sterile specimen has been referred here. San .lose: C. Zurquf, 2000-2. r .(H) m., Standi, u ,0 ./. Valerio 48292. Eulkptocium Tuck., Gen. Lich. 95. 1872. Stephmiophorus Fw., Linnaea 17: 29. 1843, Type species: no species cited. Thallus foliose, monophyllous, without rhizinae, upper and lower cortex present, each of a single layer of isodiametric cells; spores muriform, many-celled. This subgenus is predominantly tropical with many species in tropical America. Le&togium olivaceum (Hook.) Zahlbr., Cat. Lich. Univ. 3: 146. 1924, excl. syn. Collcma olivaceum Hook. ap. Kunth, Syn. PL Aequinoct. Orb. Nov. 1 : 38. 1822. Type: Colombia, Cauca, between Popayan and Almaguer, Humboldt 252. Thallus isabella to light brownish olive, lobes broad and round- ed, surface deeply reticulate-scrobiculate with sharp ridges and margins granulate-isidiose, or ridges rarely microphylline; about 200 ijl thick, corticate above and below with an algal layer of loosely Singled filaments of Nostoc corresponding to a medulla, separated from the cortex by a palisade layer about 30 y. thick. Apothecia sessile, margin verrucose, concolorous, disc darker; amphithecium thick, homogeneous with the thallus; parathecium about SO [l thick, of very densely woven hyphae; hypothecium about 25 (jl thick, very deeply staining; thecium about 100 [l tall; paraphyses filiform, about 1.5 y. in diameter, ending in the epithecial gel. The apothecia are very rare and too immature in the one specimen from Costa Rica to give all the characters, but it seems quite distinct from L. reticulatum and />. foveolalum. Cartago: It. Reventaz6n below Santiago, 710-750 m., Dodge 8010. Var. granulosum Dodge, var. n<>v. Type: Costa Rica, San Jose, H. Virilla below El Brazil, Dodge 8030. Thallus similis speciei sed nigro-granulatum ambobus super- ficiebus. Apothecia obscure brunnea, nigricantia, sessilia, basi 1933] DODGE — LICHENS OF COSTA RICA. I 425 constricta, disco concolore; amphithecium 190 ^ crassitudine, corticatum cellulis isodiametricis; parathecium 130-140 ;x crassi- tudine, ad marginem 40 [x attenuatus, hyphis dense contextum; hypothecium 30 y. crassitudine; thecium 120-130 \x altum; paraphyses similes speciei; asci 12-16 ;x diametro metientes; ascosporae imbricatim monostichae, octonae, fusiformes, 22-24 x 12-14 [i. Thallus similar to that of the species but black-granulate on both surfaces. Apothecia dark brown, blackening, sessile, constricted at the base, disc concolorous; amphithecium 190 y. thick, homogeneous with the thallus, corticate with a single layer of isodiametric cells; parathecium 130-140 tx thick, thinning toward the margin to 40 \i, of densely woven hyphae ; hypothecium 30 tx thick, so deeply staining that its structure is not clear; thecium 120-130 [x tall; paraphyses as in the species; asci cylindric, 12-16 [i in diameter; ascospores imbricately monostichous, 8 per ascus, fusiform, 22-24 x 12-14 \i. San Jos6: R. Virilla below El Brazil, Dodge 8030. Leptogium foveolatum Nyl., Syn. Meth. Lich. 1 : 124. 1858. Type: not stated, specimens from Bolivia, Weddell, and Mexico, Fr. Muller, are mentioned. Thallus mineral gray or darker, often discolored light yellowish olive or darker, lobes rounded, smooth, surface deeply foveolate with sharp wrinkles both above and below; very variable in thickness up to 250 [i, Nostoc very sparsely scattered throughout, rather denser and tending to form a palisade layer just below the surface, but this layer much thinner and less definite than in L. olivaceum. Apothecia about 1.5 mm. in diameter, constricted at the base, margin smooth, light buff, disc cinnamon rufous; amphithecium about 130 (jl thick, similar to the thallus in structure but algae much more abundant and closely tangled and cortex pseudoparenchymatous, about 50 [x thick; parathecium about 40 [jl thick below, thinning out and disappearing above, pseudo- parenchymatous with small cells; hypothecium about 80 [i thick, very deeply staining; thecium about 170 [i tall; paraphyses filiform, about 2 ja in diameter, tips clavate; asci 8-spored, cylindrical, about 12 ^ in diameter; ascospores imbricately [Vol. 20 420 ANNALS OF TIIK MISSOURI BOTANICAL GARDEN monostichous, 30-40x12-16 tx [only 20x10 tx in Costa Rican specimens, but obviously immature]. San Jos<5: R. Virilla below El Brazil, Dodge 7783. Leptooium microstictum Vainio, Dansk Hot. Ark. 4 11 : 18. 1920. Type: not stated, specimens cited, Mexico, Palenque, Liebmann 7415, 7416, 7428; Papantla, Liebmann. Thallus light glaucous blue or a little darker, lobes rounded, surface smooth but irregularly impressed, margin thin and smooth; about 225 [x thick, corticate on both surfaces with cells about 4 ^ in diameter, the filaments of Nostoc very loosely tangled, more or less parallel to the surface and very scanty in the middle. Apothecia 0.5-1.0(- 1.3) mm. in diameter, sessile, constricted at the base, margin thin, whole, eoncolorous with the thallus, disc pale to tawny; amphithecium about 00 (x thick, cortex pseudoparenchymatous, about 25 [x thick below, thinning out above to a double layer of cells, algal layer of tangled filaments of Nostoc; parathecium pseudoparenchymatous, about 40 (x thick, thinning above to about three layers of cells; hypothecium 30 [x thick, of densely tangled rather large hyphae suggesting pseudo- parenchyma but cells much smaller than in the parathecium; thecium about 100 [x tall; paraphyses filiform, 1 [x in diameter, ending in the epithecial gel; asci clavate to cylindrical, 12-16 [x in diameter; ascospores distichous, 8 per aseus, fusiform, with about 5 transverse septa, 20-30 x 8-10 [x. V single small specimen from Costa Kica is doubtfully referred here. The thallus is somewhat more finely impressed and oc- casionally near the margin it appears very minutely scrobiculate. This specimen is from a greater elevation than most other mem- bers of this subgenus. San Jon<5: ( luayabillos and Cabeza de Vaca, 2150-2350 m., Dodge & Thomas 7418. Leptogium vesiculosum (Sw.) Malme, Ark. f. Bot. 19 14-15. 1924. Lichen rcsiculosus Sw., Nov. Gen. Sp, PI. Prodr. 147. 1788. Lichen Imllatum Ach., Lichen og. Suec. Prodr. 137. 1798. Leptogium bullatum Mont., Ann. Sci. Nat. Bot. II, 16: 113. 1841. Type: Jamaica, tops of mountains, 0. Swartz. L. bullatum Ach. based on the same material. H 1933] DODGE — LICHENS OF COSTA RICA. I 427 Thallus plumbeous, lobes rounded, more or less appressed, longitudinally wrinkled with shallower cross ridges; about 100 [l thick between the ridges, corticate on each surface with a single layer of isodiametric cells, homoeomerous, of very loosely tangled filaments of Nostoc running more or less parallel to the surface. Apothecia sunk in the tips of podetiiform elevations of the thallus, margin wrinkled but not isidioid or phyllophorous, edge of parathecium conspicuous light buff, disc concave, rufous; amphithecium continuous with the thallus without differentiation, cortex a single layer of cells both above and below; parathecium 100 [x thick below, thinning out to 40-60 \i above at the margin, pseudoparenchymatous ; hypothecium thick, 80 [x, of densely woven slender hyphae; thecium 120-140 (jl tall; paraphyses slender, filiform, cells almost isodiametric above, disintegrating into the brown epithecial gel; asci cylindrical, 20 ;jl in diameter, 8-spored; ascospores imbricately monostichous, broadly fusiform, 23-32 x 10-13 [a, with 5 transverse septa. This species seems to be widespread in the American tropics, from 700 to 2100 m., rarely also below these levels. L. slipitatum, with which this species is easily confused, seems confined to the lower levels, reaching up to about 800 m., and has so far only been reported from Guadeloupe and Costa Rica. Lim6n: Waldeck, Dodge & Nevermann 7938; Marta, 20 m., Dodge 7427. Cartago: Angostura, Polakowsky 456; Juan Vinas, 1000 m., Calvert 68; Santiago, 1140-1180 m., Dodge 4555; R. Birrfs, 1220-1340 m., Dodge & Thomas 4628; Agua- caliente, 1240-1460 m., Dodge & Thomas 7082; Carpintera, 1700 m., K. Danielson 101a. San Jos6: S. Pedro de Montes de Oca, 1200 m., Thomas 4717; S. Juan Tibas, 1000-1100 m., Dodge 4303. Guanacaste: H. Santamaria, 640-680 m., Dodge & Thomas 6797. Puntarenas: Boruca, 560 m., Tonduz 5380. Var. digitatum Eschw. ap. Martius, FL Brasil. 1 : 238. 1833. Collema bullatum Raddi, Atti Soc. Ital. 18: 36, pi. 4,f. 2. Collema bullatum var. dactylinoideum NyL, Flora 41 : 338. 1858; Syn. Meth. Lich. 1: 129. 1858. Type: Brasil, between Mandiocca and Morro do Frade, Raddi, type of var. dactylinoideum not cited, but specimens from Mexico, Fr. Muller, Colombia, Tolima, Goudot, and Bolivia, Weddell, are mentioned. This variety is stated by Eschweiler to have a thicker thallus, 428 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 20 deep green instead of lead color, densely wrinkled instead of granular; podetia cylindrical instead of ventricose, 0-12 mm. instead of 2-6 mm. tall, about 6 mm. in diameter. From the description this variety does not seem distinct from the species and I have seen no material referable here from Costa Rica. It is possible that Esc h wetter was attempting to separate this species from L. phyllocarpum which he called Coll em a (Lcptogium) bullatum var. sertatum. Leftogium stiimtatum Vainio, Hedwigia 38: (255). 1899. Type: Guadeloupe Island, near Cmirbeyre, P. Duss 4$4- Thallus plumbeous, soon discolored buffy citrine to Saccardo's olive, irregularly dichotomously branched, ultimate lobes 2-7 mm. broad, then crowded and confluent, rounded, acutely and irregularly wrinkled above and below ; about 200 '^ thick between the ridges, with Nosloc filaments densely coiled and tangled near the surface, mostly parallel to the surface within, much more abundant than in L. vcsiculosum, with cortex of isodiametric cells on each surface. Apothecia <>n short thalline pustules, up to 2.5 mm. in diameter, margin thick with periclinal folds (never radial as in L. veaiculosum) , disc chestnut; amphithecium about 200 [i thick, of the same texture as the thallus; parathecium 80 (i thick, disappearing above, of large-celled pseudoparenchyma; hypothecium 80 (x thick, of more or less parallel hyphae continued false parathecium: thecium ab th tips in the brown epithecial gel; asci cylindrical, 16-20 y. in diameter; ascospores distichous, broad-fusiform with acute ends, 24-32 x 10-12 ji. SSan Josi': oafion of R. Virilla, below El Brazil and Sta. Ana, Dodge 8019. Puntarenas: Peninsula of Puntarenas, 3-5 m., Dodge 8021. Leftogium phyllocarpum (Pers. ap. Gaudich.) Mont., Ann. Sci. Nat. Bot. Ill, 10: 134. 1848. Collema phyllocarpum Pers. ap. Gaudich. in Freycinet, Voy. Uranie, Bot. 204. 1826. Collema bullalum var. sertatum Eschw. ap. Martius, Fl. Brasil. 1:239. 1833. Type: Brasil, Rio de Janeiro, Gavdichaud. Thallus plumbeous but frequently becoming black, especially 1933] DODGE — LICHENS OF COSTA RICA. I 429 in specimens which have been dried slowly or repeatedly wetted and dried after collection, lobes broad, surface deeply and acutely wrinkled, the wrinkles predominantly longitudinal or radial, the secondary wrinkles very irregular; more than 200 [x thick between the wrinkles, corticate on both surfaces by a single layer of isodiametric cells, the filaments of Nostoc loosely tangled, more or less parallel to the surface and much more coiled and densely tangled just under the upper cortex. Apothecia about 2 mm. in diameter, margin with crowded periclinal wrinkles, disc rufous, immersed in hollow protuberances as in L. vesi- culosum; amphithecium appearing lobed in section up to 500 p. thick, homogeneous with the thallus but algae a little denser; parathecium about 100 [t. thick below, thinning out to about 20 (x above at the margin, of large-celled pseudoparenchyma ; hypo- thecium about 30 (i thick, of slender densely woven hyphae continuing up to the margin at about the same width; thecium about 140 tj. tall; paraphyses filiform, not dilated above, epithecial gel brown; asci cylindric, about 20 ^ in diameter, 8-spored; ascospores broad-fusiform with acute ends, monostichous, with 5 transverse septa, 22-30 (- 35) x 10-1 2 (- 14) p.. Apparently this species is very widespread and common in the American tropics from low elevations in the southern United States to 2500 m. in Peru. In Costa Rica it ranges from 500 m. to about 1900 m., being characteristic in the moister fog-bathed areas of the temperate region. Cartago: Santiago, 1140-1180 m., Dodge 8000; R. Birris, 1220-1340 m., Dodge & Thomas 7994. San Jos6: Canon of R. Virilla below El Brazil, Dodge 7782; Zapote, 1200 m., Standley 40273; La Palma, 1500-1700 m., Maxon & Harvey 7891; Quebradillas, near Sta. Maria de Dota, 1800 m., Standley 42977; Sta. Maria de Dota, 1500-1800 m., Standley 42424; S. Marcos de Dota, 1200 m., Tonduz 5374; R. Naranjo, Tonduz 5375; S. Gabriel, Tristan 5229; S. Jose, Polakowsky 113. Alajuela: R. Ciruela, 920-980 m., Dodge & J. Valerio 4858; Santiago de S. Ramon, 1000 m., Brenes; Viento Fresco, 1600-1900 m., Standley & Torres 47771. ? Puntarenas: Corozal, 5-50 m., Dodge 8025. Var. macrocarpum Nyl., Syn. Meth. Lich. 1 : 130. 1858. Type: not cited but specimens from Aequatorial America, Humboldt, Chile, Peru, and Venezuela, hind. 1092, are mentioned. Similar to the species but apothecia large, 5-9 mm., margins lobulate, foliose. American material referred to var. daedaleum 430 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 20 Nyl. probably belongs here. Distribution in Costa Rica es- sentially similar to that of the species but not yet collected in the higher elevations. Cartago: Santiago, 1140-1180 m., Dodge 8001; R. Birrfy, 1220-1340 m., Dodge & Thomas 4682; Carpintera, 1700 in., A. Danielton 101b. San Jos6: R. Naranjo, Tonduz 6871 Alajncla: La I'alma de San Ramon, 1250 m., Brenet 91. Guanacastc: H. Santiimarfa, 640-780 in., Dodge & Thomas 6776, 8002; Tilardn, 50D -690 in., StandUy A ./. Valerio 44803, Dodge A Thomas 6569. Puntarenas: Corosal, 5-50 m., Dodge 7699. Var. campestre Malme, Ark. f. Bot. 19 s : 13. 1924. Type: Brasil, Bahia, R. Vermelho, Malme 29, 30; Matto Grosso, Cuyaba, Malme 2715c, etc.; Serra da Chapada, pr. Bocca da Serra, Malme 2245 B. Differs from the typical form by narrower crowded lobes, 1 >ed diameter, margin single laver of ce thecium 00 70 u. thick, reaching the margin above: thecium red distichous, br with 5 transvt 27 x 8-10 [x, usually San Jos,': Turrticares, 540-600 m., Dodge & Thomas 7481. Alajuela: R. Ciruela, 920-980 in., Dodge A- J. Valeria 7921; Santiago de S. Ram6n, 1000 m., limns. Leptoqium coralloideum Vainio, Ann. Acad. Sci. Fenn. A6 7 :110. 1915. Leptogium diaphanum f. coralloideum Mey. & Fw., Nova Acta Acad. Leopold. Carolin. 19: Suppl. 220. 1843. Leptogium phyllocarpum var. isidiosum Nyl., Syn. Meth. Lich. 1: 130. 1858. Leptogium phyllocarpum var. coralloideum Hue, Nouv. Arch. Museum Paris III, 10: 228. 1898. The type of var. isidiosum is from Mexico, Orizaba, Fr. Mailer. Thallus as in L. phyllocarpum but densely isidiose, especially along the wrinkles, somewhat thicker, semi-pellucid. Apothecia large when present, very rare, margin densely coralloid-isidiose, disc rufous; amphithecium highly developed, homogeneous with the thallus; parathecium 140 [a thick, disappearing above, of large thin-walled pseudoparenohyma; hypothecium 60 [x thick, reaching the margin above, of densely woven slender hyphae; 1933] DODGE — LICHENS OF COSTA RICA. I 431 um thecium 200 (jl tall; paraphyses filiform with clavate apices; asci cylindrical; ascospores 8 per ascus, imbricately monostichous, ellipsoid to fusiform, apices acute, 30-40(-45) x 12-17 [x, with 7 transverse septa. This species is perhaps only a large isidiose variety of L. phyllocarpum, essentially similar in morphology but larger in most dimensions. It occupies the same general region as L. It is usually sterile and hence not easily distin- guished from sterile L. marginellum var. isidiosellum. Material has been seen from Mexico, Peru, and Brasil. Limon: Hamburg, 20-30 m., Dodge 11*20. Cartago: Pejivalle, 650-900 m., Standley & J. Valerio, Dodge & Thomas 4387; Canon of R. Reventaz6n below Santiago, 920-1000 m., Dodge & Thomas 8013, Dodge 8004; R- Birris, 1220-1340 m., Dodge & Thomas 7954; C. Carpintera, 1320- 1700 m., Dodge 3758, Dodge & Thomas 4762, K. Danielson 98; Aguacaliente, 1240- 1460 m., Dodge & Thomas 8003; R. Reventado, 1460-1650 m., Standley & J. Valerio 49614- San Jose: R. Virilla below El Brazil, Dodge 6496; Zurquf, 1600-2500 m., Standley & J. Valerio 48255, 48266, 48146, Dodge, J. Valerio & Thomas 6042. Alajuela: near Fraijanes, 1500-1700 m., Standley & Torres 47425, 47462. Guanacaste: H. Santamarfa, 640-680 m., Dodge & Thomas 8005; Tilardn, 500-650 m., Standley & J. Valerio 44367, 44403. Leptogium marginellum (Sw.) S. F. Gray, Nat. Arrang. Brit. PL 1:401. 1821. Lichen marginellus Sw., Nov. Gen. Sp. PL Prodr. 147. 1788. Collema vesicatum Taylor, London Jour. Bot. 6: 196. 1847. Leptogium corrugatulum NyL, Syn. Meth. Lich. 1 : 132. 1858. Type: Jamaica, Blue Ridge Mt., Swartz. The type of C. vesicatum Taylor is from St. Vincents, West Indies. The type of Leptogium corrugatulum is from Mexico, Jalapa, Galeotti 9630. Thallus plumbeous, round-lobed, margin smooth or occasionally microphylline, pellucid, surface of undulating longitudinal wrinkles less acute than in L. phyllocarpum, of variable thickness, about 160 n thick, corticate on both surfaces with small isodia- metric cells. Apothecia minute, up to 0.3 mm. in diameter, margin densely isidiose, disc rufous; amphi thecium thick, with pseudo- parenchymatous cortex below, a single layer of cells above, as are also the isidia; parathecium 150 p thick, pseudoparenchymatous; thecium 150-180 \x tall, epithecium tawny; asci 8-spored; spores imbricately monostichous, ellipsoid-fusiform, ends acute, 25-35 (-^0) x 10-13 \l, with 5 transverse septa. 432 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 20 This species rarely matures spores although rudimentary apothecia are nearly always abundant. The microscopic details of the apothecium are largely taken from Malme as I have seen no main re asci in my Costa Rican material. There is consider- able variation in the amount of isidia, as they easily break off leaving minute foveoles on the margin. The species is quite distinct and is widespread in the American tropics. I have seen material from Florida, Alabama, Bermuda, various islands of the West Indies, Mexico, Brasil, and the (Jalapagos Islands. It seems to range from sea level to 1500 m. From the nature of the isidia it seems likely that var. isidio- sellum Kiddle, Brooklyn Bot. Gard. Mem. 1: 115. 1918, should be referred to L. coralloideum, although sterile isidiose states are very difficult to place. Limtin: Mart a, 20 m., Dodge & Nevermann 7986; Carmen, Dodge 7422; Waldeck, Dodge & Nevermann 7J+8L Cartago: Pejivalle, (>(>0-900 m., Dodge & Thomas 8014 y Staudley & J. Valeria 46769; It. Hints, 020-1100 m., Dodge 7988; Santiago, 1140 1180 m., Dodge 7987; Aguacaliente, 1240-1460 m. f Dodge A Thomas 7937; Carpintera, 1500 m., Dodge 3948. Alajurla: Santiago de S. Ram6n, 1000 m., Hrenes 242; S. Pedro de S. Rara6n, 700 in., H rents. Guanacaste: H. Kantamarfa, 610-720 m., Dodge & Thomas 6867, 7989; H. Grana- dilla, above R. S. Jos6, 4S0 m., Dodge & Thomas 7936; H. Q. Azul, on lower slope of V. Tenorio, 400 600 m., Dodge & Thomas 6653; Tilardn, 500-600 m., Dodge & Thomas 6561. Puntarenas: It. Terrones, 30 in., Dodge eft Marks 7980; Corozal, 5-50 m., Dodge 7533. Leptogium denticulatum NyL, Ann. Sci. Nat. Bot. V, 7: 302. 1867. L. tremelloides v. leptophyllinum Mey. & Fw., Nova Acta Acad. Leopold, Carolin. 19: Suppl. 228. 1843. L. tremelloides v. azureum f. isidiosum Mull. Arg., Bull. Soc. R. Bot, Belg. 29: 49. 1891, non Flora 65: 292. 1882. L. leptophyllinum Zahlbr., Cat, Lich. Univ. 3: 136. 1924. Type: Colombia, San Jil, 1300 in., Lindig. Thallus mineral gray or darker, large, broadly lobed, lobes round, edges smooth, somewhat crisped, surface slightly wrinkled, 100-120 [i thick, microphylline, isidiose, corticate on both sides with a single layer of cells, the short filaments of Nostoc with cells 4-5 [i in diameter mostly assembled just under the cortex, 1933] DODGE — LICHENS OF COSTA RICA. I 433 leaving the central portion relatively free of algae. Apothecia relatively rare but when present usually numerous, scattered over the upper surface, up to 2 mm. in diameter, margin granular, thick, becoming isidiose in age, buff, disc rufous; amphithecium 220 p. thick, with a layer of pseudoparenchymatous cortex below about 160 [x thick, not thinning out much above and merging with the parathecium, algal layer 60 [x thick below, thinning out to 30 \x above; parathecium 100 [x thick below, thinning out to about 40 [l above, of large thin-walled hyphae at times sug- gesting pseudoparenchyma; hypothecium 20 \x thick, of more slender deeply staining hyphae; thecium 160 pi tall; paraphyses slender, ending abruptly in the epithecial gel; asci 10-12 [x in diameter, cylindrical, 8-spored; ascospores imbricately mono- stichous, fusiform with acute ends and about 5 transverse septa, about 24 x 10 [x. This species seems widespread in the American tropics, al- though it is so rarely fertile that it is difficult to be sure of the determination of much of the sterile material, especially to separate it from L. cyanescens var. austro-americanum Malme. As here described, this species is close to, although not identical with, Malme's variety, while the L. denticulatum Malme seems to be an isidiose state of or closely related to L. Tuckermani. Only a careful comparison of all the fertile specimens from tropical America in the various herbaria of Europe can settle these problems. Nevermann Waldeck 7997, Dodge 8012. 920-1000 100-1 Dodge & Thomas 7992; Estrella, 1600 m., K. Danielson 162; near Cartago, 1500 m., K. Danielson 22; C. Carpintera, 1500 m., Dodge 397S. San Jos6: San Jose, Tonduz 5247. Heredia: C. Zurquf, 1600-1700 m., Dodge, J. Valerio & Thomas 8048. Alajuela: C. Mondongo de S. Ram6n, 750-800 m., Brenes 275; Alto de la Palma de S. Ram6n, 1250 m., Brenes 877. Guanacaste: H. Santamarfa, 680-780 m., Dodge & Thomas 6870, 7014; H. Grana- dilla above R. Las Canas, 500-600 m., Thomas 6623. Puntarenas: Osa, Corozal, 5-50 m., Dodge 7998. Leptogium azureum (Sw.) Mont. ap. Webb, Hist. Nat. lies Canaries 3 1 : 129. 1840. 434 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 20 Lichen azureum Sw. ap. Ach., Lichenog. Suec. Prodr. 137. 1798. Type: mountains of Jamaica, 0. Swartz. Thallus pellucid, greenish -glaucous-blue to puritan gray, broadly sinuate-rounded lobes, smooth; margins smooth, 80- 100 [i thick, of loosely coiled and tangled filaments of Nostoc with ellipsoidal cells 2.5x4 [x. Apothecia scattered, 1-2 mm. in diameter, margin smooth, light buff, disc rufous, sessile or on a short stalk; amphithecium 320 ^ thick, below with a pseudo- parenchyma tous cortex about 100 (x thick, above with a single layer of isodiametric cells in the cortex; parathecium filamentous, 40 [i thick, poorly developed, of slender hyphae not reaching the margin; hypothecium 20 [i. thick, deeply staining; thecium 120- 150 fj. tall; paraphyses filiform, 1.5-2 \x thick, thickened at the tips to about 4 y.; asci clavate-cylindrical, about 18 \i in diameter, 8-spored; ascospores imbricately monostichous or somewhat distichous, fusiform ends acute, with 3 or 5 transverse septa, 22-28 x 8-10 [x. Only a careful study of all the types involved can settle no- menclature of the group of species centering around L. tremel- loides. This species was originally collected in South Africa by Thunberg, and very briefly described by Linne fil. The early lichenologists of the west coast of Europe identified their material as this species and later reduced Swartz' Jamaican L. azureum to synonymy. There is apparently a whole group of species having approximately the same macroscopic appearance but wholly different microscopic structure which are at present referred to this species by various workers. As we have here defined it (agreeing closely with L. tremclloides Malme), it is a species of the lowlands coming up the river valleys under favor- able conditions to about 1000 m. in Costa Rica. It seems widely distributed at lower elevations in tropical America. Lim6n: Martii, 20 m., Dodge 573; Waldeck, Dodge & Nevtrmann 7412, 7418, 7416; Gudpiles, 300- 500 m., Standi*!/ 87137. Cartago: Tunialba, 0rded; R. Pejivalle, 650-800 m., Dodge db Thomas 4414; R. Reventazuii, 920-1000 in., Dodge & Thomas 4604. 1933] DODGE — LICHENS OF COSTA RICA. I 435 Guanacaste: H. Santamarfa, 760-900 m., Dodge & Thomas 6876; Tilardn, 500-690 m., Dodge & Thomas 8009. Puntarenaa: Osa, C. Guaca, 85 m., Dodge 7505; Corozal, 5-50 ra., Dodge 7580. Isla Coco: Snodgrass & Heller. Leptogium Standleyi Dodge, sp. nov. Leptogium tremelloides Auct., non Linne" fil. Type : Costa Rica, La Hondura, Standley 37870. Thallus plumbeus, crassus, laevis vel minute rugulosus, lobis rotundatis, latis, marginibus integris, 150-170 (x crassitudine, filamentis nostocaceis sub cortice dense implicatis, centro laxe aut sparse implicatis, corticatum e serie simplice cellularum. Apothecia ad 3 mm. latitudine, plana, margine integro, tenui, laevi, fulva, disco rufo; amphithecium inferne 160 jx, ad 100 y. crassitudine superne in margine attenuatum, cortice pseudo- parenchymatico, 90 [x crassitudine inferne ad una serie cellularum superne attenuatum, filamentis nostocaceis dense contextum; parathecium 60 {x crassitudine, hyphis tenuibus dense contextum ; hypothecium flavum, 20-30 ix crassitudine, hyphis tenuibus dense contextum; thecium ad 180 {x altitudine; paraphyses filiformes, tenues, apice non incrassata; asci cylindrici, 20 (x diametro metientes ; ascosporae octonae, imbricatim monostichae vel distichae, fusiformes, 24-28 x 12 (x. Thallus mineral gray or darker, thick, smooth or minutely and shallowly wrinkled, lobes broad, rounded, margins smooth, 150-170 [x thick, filaments of Nostoc densely tangled next the cortex, looselv taneled and scattered in the center of the thallus, of both surfaces. Apothecia 3 mm. broad, plane, margin smooth, thin, tawny, disc rufous; amphithecium 160 [x thick below, thinning to 100 [x above at the margin, corticate below, 90 ;x thick, pseudoparenchymatous, above becoming a single layer of cells, filaments of Nostoc densely tangled; parathecium 60 ;x thick, of slender more or less parallel hyphae ; hypothecium yellowish, 20-30 y. thick, of slender densely woven hyphae; thecium 180 ix tall; paraphyses filiform, slender, apices not thickened; asci cylindrical, 8-spored, 20 ^x in diameter; ascospores imbricately monostichous or distichous, fusiform, 24-28 x 12 ix. This species is essentially close to L. azureum in its microscopic characters but differs in its much larger dimensions, its thicker, 436 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 20 opaque tlmllus, and larger apothecia. Superficially it some- what resembles L. pulchdlum but is much less coarsely wrinkled and differs in microscopic details. In Costa Rica it seems to be confined to the higher elevations reaching 1700 m. It is probable that material determined as L. pulckettum from Costa Rica by M tiller Argau belongs here, although I did not have time to section it. It certainly is not typical L. pulchdlum. Cartago: Turrialba, Orxtcd; Orosf, 1000-1100 m., Standby 39791. San ,Jos«': San Jos<5, 1 130 m., Standby 41883; La Hondura, 1300-1700 m., Standlcy 87870, tv no. Alajuela: Pods, Tondiu 5801. Leptogium Tuckermani Dodge, sp. nov. Leptogium trerndloides var. minor Tuck, in herb. ? Leptogium moluccanum Vainio, Etude Lich. Br6sil 1 : 223-224. 1890, non Collema moluccanum Pers. ap. Gaudich. in Freycinet, Voy. Uranie, Bot. 203. 1826. Type: Cuba, Monte Verde, C. Wright 56. Thallus plumbeus vel obscurior, lobis confertis, rotundatis, lerneque • isidiisoue de itudine cibu Vpothecium mm. dianietro metiens, peltatum. basi margino integro, pallescente, disco castaneo; aniphithecium corticatum cellulis pseud«>parenchymaticis, 00 \x crassitudine inferne, attenuatum ad 12 [i superne, strato nostocaceo 40 [x crassitudine; parathecium pseudoparenchyniaticum tenue, 10- 12 \i crassitudine, cellulis parvis; hypothecium 20 (x crassitudine. hyphis tenuibus dense contextum; lliecium 90-120 jx altitudine; paraphyses filiformes, 2 fx dianietro, apicibus incrassatis; epi- thecium brunneum; asci cylindrici, 12 [x dianietro metientes; ascosporae octonae, imbricatim monostichae, apicibus acutis, niurales, septis transversalibus 3-5, cellulis haud numerosis, 18-22 x 10 12 u. Thallu bes crowded d crisped, rounded, smooth on both sides, 35-00 ex layer of Nostoc about 25 fx, with pseudoparenchyma- above and below of large cells. Apothecia minute, . in diameter, constricted at the base, margin whole, e, disc chestnut; aniphithecium with a thick pseudo- 1933] DODGE — LICHENS OF COSTA RICA. I 437 parenchymatous cortex below, about 60 ^jl, thinning to two rows of cells about 10-12 (x thick above; parathecium thin, pseudo- parenchymatous, 10-12 (x thick, of small cells, inconspicuous and often reported absent; hypothecium 20 ex thick, of slender densely woven hyphae; thecium 90-120 [x tall; paraphyses filiform, 2 [x in diameter, with clavate apices; epithecium brown; asci cylindric, 12 ^ in diameter; ascospores 8 per ascus, imbricately monostichous, ends acute, muriform, with 3-5 transverse septa, cells not numerous, 18-22 x 10-12 [x. This species has long been confused with L. moluccanum, L. Mariannum, and L. diaphanum. It differs from the two former in habit and color and from the latter by its structure. It is apparently widespread in the American tropics, being reported from Paraguay, and from Brasil in the states of Rio Grande do Sul and Matto Grosso by Malme, and in Minas Geraes by Vainio. Lim6n: Hamburg, Dodge 7425. Cartago: Pejivalle, 680 m., Dodge & Thomas 4556; Santiago, 1140-1180 m., Dodge 4555, 80 M; Las Concavas, 1350-1450 m., Dodge 6842. San Jos6: hills above Sta. Ana, Dodge 7781. Guanacaste: H. Santamarfa, 680-780 m., Dodge & Thomas 8043; H. Granadilla, 480 m., Dodge & Thomas 8045. Leptogium simplicius Vainio, Ann. Acad. Sci. Fenn. A6 7 : 109. 1915. Type: S. Domingo, La Cumbra, C. Raunkiaer. Thallus between light drab and light grayish olive, with irregularly incised lobes 5-7 mm. broad, not isidiose, with slightly undulate elevated acute wrinkles on both surfaces, corticate with a single layer of isodiametric cells* Apothecia broadly adnate, constricted at the base, sessile, 1.5-2.2 mm. in diameter, disc plane, rufous, margin thin, smooth, lighter than the thallus; amphi thecium 140-150 tx thick, corticate with a single series of cells above, pseudoparenchymatous below, 90-100 fx thick; parathecium 120 [jl thick below, thinning out to 20 y. thick at the margin above, pseudoparenchymatous; hypothecium fila- mentous, 30 (jl thick; thecium 150-170 (x thick; paraphyses filiform, tips somewhat thickened in the brownish epithecial gel; asci cylindric, 14-16 [x in diameter, 8-spored; ascospores imbri- cately monostichous, fusiform, thin-walled, with 5 transverse septa, 22-30x11-13 [x. 438 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 20 This species previously reported only from the type locality has yet been found only at low elevations on the Pacific slope. San Josr: Tumicares, 540-400 in., Dodge A Thomas 801+7. Guanacaste: Liberia, 100 in., Dodge, Alfaro & Thomas 6586, Dodge & Thomas 8028, Tihinin, 500 600 m., Dodge u>x of two or more rows of cells; hyphae perpendicular to the surface, medulla loosely woven. Apotheeia lecanorine. Tomentum absent or very early evanescent, algae definitely Nostoc, colonies usually separate; parathecium usually pseudoparenchj matous Pannaria Tomentum thin hut persistent, algae Nostoc ? or perhaps Scytonema; parathecium of slender hyphae Malmella Apotheeia biatorine, pseudoparenchymatous from periclinal hyphae. Tomentum absent, algae Nostoc; rhizinae well developed. .. .Parmeliella Tomentum present and highly developed, algae Scytonema or Nostoc; lower cortex little developed; apotheeia large, mar- ginal Erioderma 1933] DODGE — LICHENS OF COSTA RICA. I 441 Upper cortex of septate periclinal hyphae which may simulate pseudo- parenchyma, tomentum usually absent; apothecia biatorine, medulla of periclinal conglutinate hyphae appearing pseudoparenchymatous, algae Scytonema Coccocarpia PANNARIA Del. ap. Bory Pannaria Del. ap. Bory, Diet. Class. Hist. Nat. 13 : 20. 1828. Type species: Pannaria rubiginosa (Thunberg ap. Ach.) Del. ap. Bory. Thallus granular, squamose to foliose with a well-developed bluish black or black hypothallus, rarely with dark, more or less tangled rhizinae below, heteromerous, upper surface corticate with large-celled pseudoparenchyma formed from a palisade of hyphae; algae Nostoc, medulla single or double, in the former case arachnoid, in the latter the upper portion of more or less parallel, thin-walled hyphae, loosely woven, and below of densely tangled hyphae, without lower cortex. Apothecia at first sunk in the thallus, finally sessile or peltate, superficial ; amphithecium pseudoparenchymatous with a few algae in the center; hypo- thecium hyaline; asci clavate, 8-spored; ascospores hyaline, elongate, ellipsoidal to almost fusiform with a somewhat thick- ened and finely verrucose wall. Spermagonia in hemispherical warts, spermatiophores septate with short, broad cells; spermatia straight or very slightly curved, elongate, cylindrical. This genus seems one of the most variable and widespread of the family, occurring from the Arctic to the Antarctic, in Costa Rica occurring from sea level to 1700 m. In tropical America the genus separates easily into two sections, the isidiose species with a filamentous parathecium, and the smooth species with a pseudoparenchymatous parathecium. KEY TO THE TROPICAL AMERICAN SPECIES OF PANNARIA Hypothallus not well developed, pale with pale rhizinae below. Thecium 80-100 y. or less tall; parathecium filamentous; hypothecium not differentiated; spores 11-12x4-5 \x y more or less constricted at the middle and appearing 2-celled; S. Africa P. rubiginosa Thecium 100-120 /i or more tall; parathecium of septate hyphae; hypo- thecium pseudoparenchymatous; spores 13-16 x 7-9 ju; Rio Grande do Sul f Brasil [P. rubiginosa Malme non aliorum] P. Malmei Hypothallus developed, sometimes sparingly. Hypothallus pale, thallus squamulose, pale olivaceous to rusty yellow, margin cut-crenate, irregularly lobulate; spores 8-9 x 2.5-3 a*; Mexico. P. applanata [Vol. 20 442 ANNALS OF THE MISSOURI HOTANICAL GARDEN Hypothalliis black. Thallus leidioid. Margins and surface with cylindrical, coralloid branched isidia; spores 14-19 x 7-8.5 mJ Antilles and Brasil P. stylophora Margins isidio-lacinulate; spores 8-10 x 5-6 /x, with thick epispore; Minas Geraes, Brasil P. isidiovlra Margins with verruciform isidia, lobes under 1 mm. broad; apothecia 1-1.5 mm. in diam.; spores 12-14 x 8-10 m, wall thin and smooth. P. Moseni Thallus without isidia or soredia. Thallus livid fusceeoent, mdiately broad-lobed; spores 14 x 6/z; hypo- thallus less developed, center glebulose-lobed; Sao Paulo, Brasil. P. imbricatula Thallus pallid or ashy or pale glaucous, lobes subcuneate, up to 2 mm. broad; spores 1 2— 1 5 (— 1 S ) xS 9(-10) a*; parathecium and hypotheciuni pseudoparenchymalous P. Vainii Thallus coeruleo-ashy, lobes cuneate obovate; epilhecium black aeruginoue; spores 12-20 x 7-8 /z, surface subplicate; Sao Paulo, Brasil P. caeruleo-nigricans Thallus livid glaucescent or lurid pallid, laciniae very distinctly radiate-stellate, even subimbricate P. radiata Thallus pale fuscescent, lobes 2-2.5 mm. broad and about 6 mm. long; epithecium rufous; spores 12 19x8-10 m; parathecium p«eudoparenchymatous; hypotheciuni of conglutinate vertical hvphae; Brasil P. brasiliensis Pannaria isidioidea Vainio, Ann. Acad. Sci. Fenn. A6 7 : 102. 1915. Pannaria Mariana var. isidioidea Vainio, Etude Lich. Brasil 1:200. 1890, excl. syn. Type: lirasil, Minas Geraes, Sitio, 1000 m., Vainio 669, 98,! (sterile). The following description is based on fertile specimens from Costa Rica. Hypothallus of dark green rhizinae extending only slightly beyond the thallus; thallus dark olive buff, margins lighter, surface very minutely tomentose, pinnatifid dissected, lobes somewhat cuneate, about 1 mm. wide, ultimate lobules rounded, 0.25-0.5 mm. broad, toward center becoming microphylline and subisidiose but isidia somewhat flattened, rarely cylindric, 150- 160 [a thick, upper 40 \l of pseudoparenchymatous cortex, algal layer of Nostoc about 40 \i thick, and medulla 40 \i thick, of loosely woven hyphae with occasional small chains of Nostoc; rhizinae thick-walled, black, 0-8 \i in diameter. Apothecia crowded, peltate, constricted at the base, about 1 mm. in di- ameter, margin prominent, crenulate, almost lobulate, disc 1933] DODGE — LICHENS OF COSTA RICA. I 443 chestnut; amphithecium 200 \l thick, of the same texture as the cortex and algal layer of the thallus; parathecium 60 y. thick, filamentous, of large thin-walled periclinal hyphae, ending above in a false pseudoparenchyma ; hypothecium about 20 y. thick, of similar structure but more deeply staining; thecium 80-100 |x tall; paraphyses slender, filiform, somewhat clavate above in the brownish epithecial gel; asci clavate, 12 (x in diameter; ascospores ellipsoidal with a thick epispore, 8-10 x 5-6 y. (perhaps still immature) . This species is somewhat variable in appearance and perhaps should be separated into varieties and forms but I have not seen sufficient material to do so properly. Sterile material has been referred here solely on the structure of the thallus. Dodge & Thomas 6558 from Tilaran has a white reticulate cortex. Cartago: R. Birds, 1220-1340 m., Dodge & Thomas 4559. Heredia: C. Central de Zurquf, 1600-1700 m., Dodge, J. Valerio & Thomas 6048. Alajuela: Piedades de S. Ram6n, 900 m., Brenes S75. Guanacaste: H. Santamarfa, 640-680 m., Dodge & Thomas 682S, 6911, 6988; Tilaran, 500-690 m., Dodge & Thomas 8059, 6558. Var. pulvinata Dodge, var. nov. Type: Costa Rica, Alajuela, C. de Pata de Gallo a S. Rafael de S. Ram6n, 1200-1250 m., Brenes. Thallus ut in P. isidioidea sed isidiosissimus, marginibus adscendentibus, cinerascentibus ; isidia coralloidea, elongata, pulvinum formantia. Thallus as in P. isidioidea but densely isidiose with ascending, cinerascent margins; the isidia coralloid, repeatedly branched, crowded, forming a cushion up to 2 cm. in diameter and 3-4 mm. thick. This variety might easily be mistaken for a sterile pulvinate Stereocaulon or a Siphula, were it not for the occasional lobe at show 1200-1250 Pannaria Moseni Dodge, sp. nov. Pannaria rubiginosa Malme, Ark. f. Bot. 20 3 : 7-8. 1924, pro parte. Type: Brasil, Sao Paulo, Sororocaba near Santos, Mosen 8284. Thallus albidus, subpinnatifidus, lobis 3-5 mm. longitudine, 0.8 mm. latitudine, apicibus rotundatis, marginibus loborum 444 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 20 verrucosis, vel etiam in centro thalli isidiosus, ad 140 [x crassitu- dine, cortex superior 30-35 ^crassitudine, pseudoparenchymaticus ex hyphie perpendicularibua formatus, zona nostocacea coloniis Bubsphericis 35 [x diametro metientibus, medulla ad GO (x crassi- tudine hyphis laxe implexis, rhizinae brunneo-nigricantes, hyphis nigris pachydermaticis 3-4 [x diametro, conglutinatis. Apothecia 1-1.5 mm. diametro, marginibus tenuibus, rrenato-lobatis et subisidiosis; amphilhecium inferne 120 ex crassitudine superne ad 60 [l attenuatum, cortice 40 [x crassitudine, pseudoparenchy- matico; parathecium bene evolutum, inferne 40 [x superne ad 60 [i crassitudine, hyphis ad 4 \i diametro leptodermaticis con- textum; hypothecium 25 [x crassitudine, liypliis periclinalibus dense compactum; thecium ad 100 [x altitudine; paraphyses filiformes 1-2 [x diametro apicibus clavatis; asci clavati, 25 [x diametro; ascosporae octonae, distichae, 12-14x8-10 [x, epi- sporio tenuiusculo. Thallus pearl gray, subpinnatifid, lobes 3-5 x 0.8 mm., with rounded tips, margins of lobes verrucose or even occasionally cylindric, isidiose toward the center, about 140 \i thick, upper cortex 30-35 [x, pseudoparenchymatous from the breaking up of a palisade layer, algal zone of subspherical colonies of Nostoc about 35 [x in diameter, medulla about 60 [x thick, of loosely woven hyphae; rhizinae of brownish black thick-walled conglu- tinate hyphae 3-4 [x in diameter. Apothecia 1-1.5 mm. in diameter, margins thin, crenate-lobulate or even somewhat isidiose; amphithecium about 120 [x thick below, thinning to about 60 [i above with a cortex about 40 [x thick of pseudoparen- chyma; parathecium well developed, about 40 [x below, spreading to 60 [i above, of interwoven thin-walled hyphae about 4 [x in diameter; hypothecium about 25 [x thick of dense periclinal hyphae, staining very deeply; thecium about 100 jx tall; para- physes filiform, 1-2 [x in diameter, with clavate tips; asci clavate, 25 [x in diameter, 8-spored; ascospores distichous, 12-14 x 8-10 [x, with a somewhat thinner epispore. It is with some hesitation that I have described this species as new. The thallus is very suggestive of the sterile P. stylophora, although it has a somewhat different color and lobing and rather larger dimensions of the parts. It does not seem to be closely 1933] DODGE — LICHENS OF COSTA RICA. I 445 related to P. ruhiginosa, originally described from Thunberg's South African collections, and is certainly not closely related to the material referred to that species in the North Temperate zone. So far I have only seen this species from the Atlantic Coastal Plain of Costa Rica. Lim6n: Hamburg, Standley